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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY OF SCIENCES 



Vol. 27 December, 1964 No. 4 



VOLUME 27 



Editor 
Pierce Brodkorb 



Published by the 

Florida Academy of Sciences 

Gainesville, Florida 

1964 



Mailing Dates of Volume 27 



Number 1 
Number 2 
Number 3 
Number 4 



April 27, 1964 
July 14, 1964 
November 4, 1964 
February 26, 1965 



New Taxa Proposed in Volume 27 

\Calappilia brooksi Ross and Scolaro (Decapoda: Calappidae) 
\Calappa robertsi Ross, Lewis, and Scolaro (Decapoda: Calappidae) 



187 



f Aparnocondylus ocalanus Ross, Lewis, and Scolaro 

(Decapoda: Calappidae) 193 



\Balanus tamiamiensis Ross (Cirripedia: Balanidae) 
Protoneura viridis Westfall (Odonata: Protoneuridae) 



272 
112 



Sphaerodactylus copei enochrus Schwartz and Thomas 

(Reptilia: Gekkonidae) 322 

Sphaerodactylus copei cataplexis Schwartz and Thomas 

(Reptilia: Gekkonidae) 326 

Leiocephalus stictigaster parasphex Schwartz (Reptilia: Iguanidae) 212 

Leiocephalus stictigaster ophiplacodes Schwartz (Reptilia: Iguanidae) 217 

Ophisaurus ceroni Holman (Reptilia: Anguidae) 311 

\Nettion greeni Brodkorb (Aves: Anatidae) 55 

\Agriocharis progenes Brodkorb (Aves: Meleagrididae) 223 

\Fulica shufeldti Brodkorb (Aves: Rallidae) 186 



Fossil 



CONTENTS OF VOLUME 27 



Number 1 

A valuable old collection of Florida marine algae Wm. Randolph Taylor 1 

Effect of reduced water temperature on fishes of Tampa Bay, Florida 

Gordon R. Rinckey and Carl H. Saloman 9 

Western Atlantic serranid fishes (groupers) of the genus Epinephelus 

Luis Rene Rivas 17 

Fishes of some South Carolina coastal plain streams 

William D. Anderson, Jr. 31 

A Pliocene teal from South Dakota Pierce Rrodkorh 55 

Tapirus copei in the Pleistocene of Florida Clayton E. Ray 59 

Notes on the Odonata of Cuba Minter J. Westfall, Jr. 67 

Dietary vitamin A and copper concentration in the liver and heart of rats 

Marie L. Kraemer, M C. Jayaswal, J. F. Easley, and R. L. Shirley 86 

Temperatures of three breeds of yearling steers in South Florida 

C. E. Haines and M. Roger 91 

Medalists of the Florida Academy of Sciences 96 



Number 2 

A new crab from the Eocene of Florida Arnold Ross and R. J. Scolaro 97 

White-cedar stands in northern Florida 

E. A. Collins, C. D. Monk, and R. H. Spielman 107 

A new damselfly from the West Indies (Odonata: Protoneuridae) 

Minter J. Westfall, Jr. Ill 

Oxygen depletion in a Florida lake George K. Reid 120 

Frogs introduced on islands Wilfred T. Neill 127 

Ethoxyquin and vitamin E studies in poultry 

R. H. Harms, C. R. Douglas, and P. W. Waldroup 131 

Soil survey for planning urban development Victor W. Carlisle 139 

Demography of a Floridian alcoholic sample James H. Williams 148 

The shrimp Trachypeneus similis in Tampa Bay Carl H. Saloman 160 

Officers and members of the Academy 165 

iii 



Number 3 

Surface calcium-chlorinity relationships 
A new name for Fulica minor Shufeldt 



Billie Z. May 111 
Pierce Brodkorb 186 



New Eocene decapods from Florida 

Arnold Ross, Jackson E. Lewis, and R. J. Scolaro 187 



Survival potential of piranhas in Florida 
New subspecies of Leiocephalus from Cuba 
Notes on fossil turkeys 
Osteology of gallinaceous birds 



Martin A. Moe, Jr. 197 

Albert Schwartz 211 

Pierce Brodkorb 223 

/. Alan Holman 230 



NUMBER 4 

Stability of ketones to alkyl borates 

An entropy function 

A new barnacle from the Tamiami Miocene 

Type locality of Platylepas wilsoni Ross 

Glucose nutrition and longevity in oysters 

Larry Gillespie, R. M. Ingle, and Walter K. Havens- 
Notes on postlarvae of Panulirus argus 

Roy Witham, Robert M. Ingle, and Harold W. Sims, Jr. 



Carl H. Snyder 


253 


Vernon E. Derr 


255 


Arnold Ross 


271 


Arnold Ross 


•278 



Two dragonflies new to Florida 



Dennis R. Paulson 



New Atlantic coast ranges for fishes 

William D. Anderson, Jr., and Elmer J. Gutherz 

Hypertensive effect of Latrodectus venoms 

John D. McCrone and Roger J. Porter 



A new glass lizard from Veracruz, Mexico 



/. Alan Holman 



Subspeciation in Sphaerodactylus copei 

Albert Schwartz and Richard Thomas 



Isolating mechanisms in snakes 
Large quahog clams from Boca Ciega Bay 

iv 



Wilfred T. Neill 
Harold W. Sims, Jr. 



279 

289 
298 

299 

307 
311 

316 
333 
348 



Quarterly Journal 

of the 

Florida Academy of Sciences 

Vol. 27 March, 1964 No. 1 

CONTENTS 

A valuable old collection of Florida marine algae Wm. Randolph Taylor 1 

Effect of reduced water temperature on fishes of Tampa Bay, Florida 

Gordon R. Rinckey and Carl H. Saloman 9 

Western Atlantic serranid fishes (groupers) of the genus Epinephelus 

Luis Rene Rivas 17 

Fishes of some South Carolina coastal plain streams 

William D. Anderson, Jr. 31 

A Pliocene teal from South Dakota Pierce Brodkorb 55 

Tapirus copei in the Pleistocene of Florida Clayton E. Ray 59 

Notes on the Odonata of Cuba Minter J. Westfall, Jr. 67 

Dietary vitamin A and copper concentration in the fiver and heart of rats 

Marie L. Kraemer, M. C. Jayaswal, J. F. Easley, and R. L. Shirley 86 

Temperatures of three breeds of yearling steers in South Florida 

C. E. Haines and M. Roger 91 

Medalists of the Florida Academy of Sciences 96 



Mailed April 27, 1964 




Quarterly Journal of tees Florida Academy of Sciences 
Editor: Pierce Brodkorb 



The Quarterly Journal welcomes original articles containing 
significant new knowledge, or new interpretation of knowledge, in 
any field of Science. Articles must not duplicate in any substantial 
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Published by the Florida Academy of Sciences 

Printed by the Pepper Printing Company 

Gainesville, Florida 



QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY of SCIENCES 



Vol. 27 March, 1964 No. 1 



A VALUABLE OLD COLLECTION OF FLORIDA 
MARINE ALGAE 

Wm. Randolph Taylor 

A collection of several hundred mounts of marine algae which 
had been made by Mrs. Mary S. Snyder between 1890-1898, or 
which had been received by her from correspondents, laid in stor- 
age for years at the Scripps Institute of Oceanography, La Jolla, 
California. It represented material from both coasts, but when 
it was decided to discard the Atlantic Coast portion this was offered 
to me for restudy by Dr. Ralph A. Lewin on behalf of the authori- 
ties at the Institute. A representative set has been deposited in 
the herbarium of the University of Michigan and the large re- 
mainder transferred as instructed to the University of California 
at Berkeley. In many cases the specimens carried no locality or 
other data, but a substantial proportion did so, and were of ex- 
cellent quality. The study of these specimens occupied part of 
my attention while the recipient of funds from the Rackham Grad- 
uate School of the University of Michigan based on a National 
Science Foundation Institutional grant. 

The first part of the collection consists of material from New 
England. The second contains Florida algae, the basis of the 
present note, and with them Bahamian and Bermudian material 
obtained from correspondents. Such West Coast material as was 
in the collection was retained in California. In the Florida ma- 
terial the preponderance of Rhodophyceae is marked, constituting 
much more than half of the variety, and even more of the station 
records. It is possible that in the course of time some packets have 
been misplaced or lost, for it is odd that no Anadyomene, no Udo- 
teas, no pelagic Sargassa, no Galaxauras or Grateloupias are in- 



IKSIITUTtOI ■""' * 



2 Quarterly Journal of the Florida Academy of Sciences 

eluded. Identification of soft filamentous species was often im- 
practicable, since the mounts were made on very porous card, and 
it generally proved impossible to detach and restore them to an 
identifiable condition. Quite a number of potential records in 
such groups as the Ectocarpoideae and Callithamnioideae have 
thereby been lost. In general the original determinations were 
appropriate to the then prevailing state of knowledge of the algae. 
If the records had been published at the turn of the century they 
would have constituted a major contribution to our knowledge of 
the Florida marine flora. With the identifications corrected and 
the nomenclature modernized the list still offers a useful record 
of the marine flora at stations some of which may now have been 
modified by urbanization. 

Little information seems to be available regarding Mrs. Sny- 
der's phycological activities. She appears to have collected in 
New England about 1890 and 1895 and in Florida between 1892 
and 1895, when her West Coast activities began. Many Florida 
botanists endeavored to trace her visits there for me, without avail. 
For their efforts I am most grateful. She seems to have been a 
seasonal visitor rather than a resident. A little Florida material 
was received from such fellow-contributors to the Phycotheca as 
Mrs. G. A. Hall and Miss C. Messina. The name of Mrs. Floretta 
C. Curtiss, Florida's most indefatigable phycologist (Curtiss, A. H., 
1899) does not appear in the collection because by this period Mrs. 
Curtiss had become inactive. Mrs. Snyder was an active corre- 
spondent of Frank S. Collins of Maiden, Mass., and specimens of 
his collecting appear among her reliquiae. 

If one examines the Phycotheca Boreali-Americanu issued by 
Collins, Holden, and Setchell one finds in many early fascicles 
specimens contributed by Mrs. E. Snyder. Professor I. Mackenzie 
Lamb of Harvard University called to my attention fascicle 37, no. 
1850, attributed to Mrs. E. Snyder on the label but to Mrs. M. S. 
Snyder (by exclusion) on the title-page. In the herbarium of 
the New York Botanical Garden I was shown a duplicate of 
Spermothamnion snyderae Farl. marked "E. & M. S. S.". These 
last two clues led to Dean Edward Snyder, Professor of German 
at the University of Illinois, who married Mary Stoddard Patchen 
and, on retirement, moved to Pacific Beach, a section of San Diego. 
The Pacific Coast specimens contributed by Mrs. Snyder are 



Taylor: Florida Marine Algae 3 

almost all from Pacific Beach, or from La Jolla just north of San 
Diego, and were made between 1894 and 1898. Following the cus- 
tom of the time the Phycotheca specimens were attributed to her 
as the wife of Dean Snyder, i.e. Mrs. E. Snyder, except for those 
in fascicles 37 and 46, issued perhaps after his death. None of 
her East Coast specimens seem to have found their way into the 
Phycotheca. Prof. G. Neville Jones kindly searched the University 
of Illinois herbarium for Florida material of Mrs. Snyder's collect- 
ing and found almost none, though some Pacific Coast material 
was present. 

In the following list the species are cited in the sequence 
adopted for my general work on warm-water algae (Taylor, 1960). 
In this one may find cited publications on Florida algae which 
had appeared previous to the time the book went to press. In 
the present paper stations are listed in the following sequence: 
Duval Co.: St. Johns River, Mayport. St. Johns Co.: St. Augustine, 
Anastasia I., Matanzas River Inlet. Brevard Co.: Rockledge, Cape 
Malabar. St. Lucie Co.: Indian River Inlet, Jensen. Palm Beach 
Co.: Jupiter and Jupiter Inlet, Juno Ocean Beach (in Palm Beach), 
Lake Worth. Monroe Co.: Key West, Dry Tortugas Ids. Pinellas 
Co.: Clearwater and Clearwater Harbor. Hillsborough Co.: Port 
Tampa. Collier Co.: Naples. Two records from St. Simons I., 
Georgia, also appear. 

CHLOROPHYCEAE 
Ulvales 
Ulvaceae 

Enteromorpha clathrata (Roth) J. Ag.: Lake Worth. E. ramulosa (J. E. Sm.) 
Hook.; Anastasia I. E. lingulata J. Ag.: St. Johns R. E. jiexuosa 
(Wulf.) J. Ag.: St. Augustine. E. linza (L.) J. Ag.: St. Augustine, 
Anastasia I., Key West. 

Monostroma oxyspermum (Kutz.) Doty: St. Augustine. 

Ulva lactuca L.: St. Augustine, Anastasia I., Lake Worth, Key West. 

Cladophorales 
Cladophoraceae 

Chaetomorpha brachygona Harv.: Key West. 

Cladophora fuliginosa Kutz.: Key West. C. fascicularis (Mert.) Kutz.: St. 
Augustine, Matanzas R., Rockledge, Lake Worth, Key West, Dry Tor- 
tugas. 



4 Quarterly Journal of the Florida Academy of Sciences 

SlPHONACLADIALES 

Dasycladaceae 

Batophora oerstedi J. Ag.: Lake Worth. 
Cymopolia barbata (L.) Lamour.: Key West. 
Acetabularia crenuluta Lamour.: Lake Worth, Key West. 

Valoniaceae 
Dictyosphaeria cavernosa (Forssk.) B0rg.: Key West. 

SlPHONALES 

Bryopsidaceae 

Bryopsis plumosa (Huds.) C. Ag.: St. Augustine, Anastasia I., Key West. 
B. halliae Taylor: Key West. 

Caulerpaceae 

Caulerpa prolifera (Forssk.) Lamour.: Lake Worth. C. mexicana (Sond.) 
J. Ag.: Lake Worth. C. ashmeadii Harv.: Naples. C. sertuluri- 
oides (Forssk.) J. Ag.: Lake Worth. C. cupressoides (West) C. Ag.: 
Key West. C. paspaloides (Bory) Grev.: Key West. C. racemosa 
(Forssk.) J. Ag., v. laetevirens (Mont.) Weber-van Bosse: Lake Worth. 

Codiaceae 

Penicillus capitatus Lamarck: Lake Worth. 
Halimeda tuna (Ell. & Sol.) Lamour.: Lake Worth. 

Codium decorticatum (Woodw.) Howe: St. Augustine. C. taylori Silva: St. 
Augustine. 

PHAEOPHYCEAE 

ECTOCARPALES 

Ectocarpaceae 

Ectocarpus siliculosus (Dillw.) Lyngb.: St. Augustine. E. confervoides (Roth) 

Le Jolis: St. Augustine. 
Giffordia mitchellae (Harv.) Hamel?: St. Augustine. G. duchassaingianus 

(Grun.) Taylor?: Dry Tortugas. 

DlCTYOTALES 

Dictyotaceae 

Dilophus guineensis (Kiitz.) J. Ag.: Dry Tortugas. 

Dictyota dichotoma (Huds.) Lamour.: St. Augustine. D. bartayresii Lamour.: 

Lake Worth. D. ciliolata Kiitz.: Lake Worth, Dry Tortugas. D. 

dentata Lamour.: Dry Tortugas. 
Pocockiella variegata (Lamour.) Papenf.: St. Augustine. 
Stypopodium zonule (Lamour.) Papenf.: Dry Tortugas. 



Taylor: Florida Marine Algae 5 

Padina vickersiae Hoyt: Lake Worth, Key West. P. sanctae-crucis B0rg.: 
Dry Tortugas. P. gijmnospora (Kiitz.) Vick.: St. Augustine, Anastasia 
I., Dry Tortugas. 

Chordariales 
Chordariaceae 
Eudesme zosterae (J. Ag.) Kylin: Dry Tortugas. 

PUNCTARIALES 

Punctariaceae 

Petalonia fascia (O. F. Miiller) Kuntze: Anastasia I. 

Colpomenia sinuosa (Roth) Derb. & Sol.: Lake Worth, Dry Tortugas. 

Hydroclathrus clathratus (Bory) Howe: Dry Tortugas. 

Fucales 

Sargassaceae 

Sargassum filipendula C. Ag.: Key West. S. pteropleuron Grun.: Lake 
Worth. S. ramifolium Kiitz.: Lake Worth. 

RHODOPHYCEAE 

Bangiales 

Bangiaceae 

Porphyra leucosticta Thuret: St. Augustine; Anastasia I. P. umbilicalis (L.) 
J. Ag.: St. Augustine. 

Compsopogonaceae 
Compsopogon caeruleus (Balbis) Mont.: St. Johns R., Jupiter. 

Nemalionales 

Helminthocladiaceae 

Liagora farinosa (Lamour.) Howe: Dry Tortugas. L. ceranoides Lamour.: 
Dry Tortugas. L. valida Harv.: Dry Tortugas. L. mucosa Howe.: 
Key West, Dry Tortugas. L. pinnata Harv.: Dry Tortugas. 

Chaetangiaceae 

Scinaia complanata (Coll.) Cott., v. complanata: Indian River Inlet (Mrs. 
G. A. Hall). S. complanata v. intermedia B0rg.: Dry Tortugas. 

Gelidiales 

Gelidiaceae 

Gelidium crinale (Turn.) Lamour.: St. Augustine, Anastasia I. 
Pterocladia americana Taylor: Naples, with Gigartina sp. 



6 Quarterly Journal of the Florida Academy of Sciences 

Cryptonemiales 

Dumontiaceae 

Dudresnaya crassa Howe: Key West. 

Acrosymphyton caribaeum (J. Ag.) Sjost: Dry Tortugas, Key West. 

Grateloupiaceae 

Halymenia floresia (Clem.) C. Ag.: Atlantic, Dry Tortugas, Clearwater Harbor 
H. bermudensis Coll. & Howe: Cape Malabar, Key West. 

Kallymeniaceae 
Kallymenia perforata J. Ag.: Atlantic, Jupiter Inlet, Lake Worth. 

GlGARTINALES 

Gracilariaceae 

Gracilaria verrucosa (Huds.) Papenf.: St. Augustine. G. debilis (Forssk.) 
B0rg.: Key West. G. ferox J. Ag.: Key West, Dry Tortugas. G. 
foliifera (Forssk.) B0rg., v. foliifera: St. Augustine. G. foliifera v. 
angustissima (Harv.) Taylor: St. Augustine, Naples. G. mamillaris 
(Mont.) Howe: Cape Malabar. G. blodgettii Harv.: Naples. 

Solieriaceae 

Agardhiella tenera (J. Ag.) Schm.: St. Augustine, Jensen, Lake Worth, Key 
West, Naples. A. ramosissima (Harv.) Kylin: St. Augustine, Dry Tor- 
tugas. 

Eucheuma schrammii (Crouan) J. Ag.: Dry Tortugas. E. acanthocladum 

(Harv.) J. Ag.: Dry Tortugas. E. isiforme (C. Ag.) J. Ag.: Juno 
Ocean Beach, Key West. 

Cystoclonium purpureum (Huds.) Batt: Anastasia. 

Hypneaceae 

Hypnea cervicornis J. Ag.: Lake Worth. H. comuta (Lamour.) J. Ag.: Dry 
Tortugas. H. musciformis (Wulf.) Lamour.: St. Augustine, Anastasia, 
Lake Worth, Key West, Naples. 

Phyllophoraceae 
Gymnogongrus griffithsiae (Turn.) Mart.: Key West. 

Rhodymeniales 
Rhodymeniaceae 

Chrysymenia enteromorpha Harv.: Key West. 

Botryocladia occidentalis (B0rg.) Kylin: Cape Malabar, Lake Worth. 



Taylor: Florida Marine Algae 7 

Champiaceae 

Lomentaria baileyana (Harv.) Farl.: St. Augustine. L. baileyana v. filiformis 
(Harv.) Farl: Naples. L. baileyana v. siliquiformis Taylor: St. Au- 
gustine. 

Champia parvula (C. Ag.) Harv.: Key West, Naples. C. salicornioides Harv.: 
Key West. 

Ceramiales 

Ceramiaceae 

Crouania attenuata (Bonnem.) J. Ag.: Key West. 

Wrangelia argus Mont.: Lake Worth. W. bicuspidata B0rg.: Key West. 
W. penicillata C. Ag.: Key West. 

Pleonosporium borreri (J. E. Sm.) Nag.: St. Augustine, Anastasia I. 

Spermothamnion gymnocarpum Howe: Key West. 

Ceramium fastigiatum (Roth) Harv., f. fastigiatum: Naples. C. fastigiatum 
f. jlaccida H. E. Petersen: Naples. C. brevizonatum Peters., v. carai- 
bica Peters.: Lake Worth, Key West, Dry Tortugas. C. subtile J. Ag.: 
Lake Worth: C. byssoideum Harv.: St. Augustine. C. floridanum 
J. Ag.: St. Augustine, Anastasia I., Lake Worth. C. diaphanum 
(Lightf.) Roth: Anastasia I., Rockledge, Lake Worth. 

Centroceras clavulatum (C. Ag.) Mont.: Lake Worth. 

Spyridia filamentosa (Wulf.) Harv.: Lake Worth. S. clavata Kiitz.: St. Au- 
gustine, Anastasia I. S. aculeata (Schimp.) Kiitz., v. disticha B0rg.: 
Naples. 

Delesseriaceae 

Caloglossa leprieurii (Mont.) J. Ag.: St. Simons I., Georgia; St. Augustine, 

Anastasia I. 
Hypoglossum involvens (Harv.) J. Ag.: Key West. H. tenuifolium (Harv.) 

J. Ag.: St. Augustine. 

Dasyaceae 

Dasya rigidula (Kiitz.) Ardiss.: Key West. D. collinsiana Howe: Key West. 
D. corymbifera J. Ag.: Cape Malabar (sensu P.B.-A. 1745). D. ramosis- 
sima Harv.: Key West. D. harveyi Ashm.: Key West. D. pedicel- 
lata (C. Ag.) C. Ag.: Key West, Dry Tortugas. D. mollis Harv.: 
Key West. 

Heterosiphonia wurdemanni (Bail.) Falk., v. laxa B0rg.: Key West. H. gib- 
besii (Harv.) Falk.: Lake Worth, Key West. 

Rhodomelaceae 

Polysiphonia subtilissima Mont.: St. Augustine. P. havanensis Mont.: Rock- 
ledge. P. binneyi Harv.?: St. Augustine. P. denudata (Dillw.) Kiitz.: 
St. Augustine, Anastasia, Lake Worth. P. ramentacea Harv.: Key 
West. 

Bryocladia cuspidata (J. Ag.) De Toni: Silver Spring (position uncertain). 

Bryothamnion seaforthii (Turn.) Kiitz.: Cape Malabar, Lake Worth. B. tri- 



8 Quarterly Journal of the Florida Academy of Sciences 

quetrum (Gmel.) Howe: Lake Worth, Key West. 

Digenia simplex (Wulf.) C. Ag.: Key West. 

Brongniartella mucronata (Harv.) Schm.: Key West. 

Wrightiella blodgettii (Harv.) Schm.: Key West. W. tumanowiczii (Gatty) 
Schm.: Key West. 

Bostrychia radicans Mont.: St. Augustine. B. rivularis Harv.: St. Simons I., 
Georgia; St. Augustine. 

Herposiphonia tenella (C. Ag.) Ambr.: St. Augustine. 

Chondria littoralis Harv.: Key West. C. atropurpurea Harv.: Key West. 
C. tenuissima (Good. & Woodw.) C. Ag.: Lake Worth, Key West, 
Naples. C. baileyana (Mont.) Harv.: Dry Tortugas. C. cnicophylla 
(Melv.) De Toni: Lake Worth. C. dasyphila (Woodw.) C. Ag.?: 
Lake Worth. C. floridana (Coll.) Howe: Naples. C. curvilineata 
Coll. & Herv.?: Anastasia I. 

Acanthophora muscoides (L.) Bory: Port Tampa, Naples. A. spicifera (Vahl) 
B0rg.: Lake Worth, Key West. 

Laurencia corallopsis (Mont.) Howe: Key West. L. papillosa (Forssk.) Grev.: 
Lake Worth. L. gemmifera Harv.: Lake Worth, Key West, Dry Tor- 
tugas. L. obtusa (Huds.) Lamour.: Key West, Dry Tortugas. L. in- 
tricata Lamour.: Lake W 7 orth, Key West, Clearwater Harbor. L. mi- 
crocladia Kiitz.: Key West. 

Literature Cited 

Curtiss, A. H. 1899. Mrs. Floretta A. Curtis, a biographical sketch by her 
son. Jacksonville, Florida, 14 + iv pp., 3 figs., 4 pis. 

Taylor, Wm. Randolph. 1960. Marine algae of the eastern tropical and 
subtropical coasts of the Americas. University of Michigan Press, Ann 
Arbor, xi + 870 pp., 14 text-photos, 80 pis. 

Department of Botany, University of Michigan, Ann Arbor, 
Michigan. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



EFFECT OF REDUCED WATER TEMPERATURE ON 
FISHES OF TAMPA BAY, FLORIDA 

Gordon R. Rinckey and Carl H. Saloman 

Estuaries connected with the Gulf of Mexico are characterized 
by extensive shallow water areas which at times are subjected to 
rapid temperature fluctuations. The reaction of marine organisms 
to environmental changes is of importance to biologists engaged 
in estuarine ecological research. Instances of massive fish mor- 
tality, especially among sublittoral species, have resulted from a 
marked lowering of water temperature. Gunter (1941) and Gunter 
and Hildebrand (1951) reported such occurrences on the Texas 
coast. Fish kills attributed to rapid temperature reduction have 
also been recorded from the Florida Gulf coast by Storey and 
Gudger (1936) and Springer and Woodburn (1960). 

During the evening of December 12, 1962, northwest winds 
of 20-30 mph. and a low air temperature of 18.3 °F were recorded 
in the Tampa Bay area. This was the coldest temperature reported 
since the initiation of records in 1890. These severe conditions 
made it possible to observe the effects of rapid temperature de- 
pression on fish in Tampa Bay. 

Two major fish kills were observed. The first occurred on the 
morning of December 13 in Old Tampa Bay below the dam of 
Bass Lake, a freshwater impoundment (fig. 1). This site is a 
brackish-water, mud-bottom habitat having an annual salinity 
range of 4 to 28 o/oo. Water depth varies from two to four feet. 
On the following morning, December 14, another kill was noted 
in Boca Ciega Bay, north of Pass-a-Grille channel (fig. 1). These 
waters range in salinity from 28 to 36 o/oo and in depth from 4 
to 12 feet. There are numerous channels and fills in the area. 
Water temperatures at the two locations are similar. 

Orservations 

Water temperature in Old Tampa Bay ranged from 14.3 to 
15.0 °C on December 10. During the fish kill of December 13, 
a surface water temperature of 9.6° C and a salinity of 14.42 o/oo 
were recorded at 9 a.m. The bottom was completely covered with 
fish in an area 50 feet square at a depth of two feet. Most of these 



Quarterly Journal of the Florida Academy of Sciences 



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]2 Quarterly Journal of the Florida Academy of Sciences 



28° 

00 T 




Fig. 1. Stations sampled during fish kills on December 13-14, 1962, in 
Tampa Bay, Florida. 



Rinckey and Saloman: Cold and Marine Fishes 13 

fish were dead, and those still alive exhibited a loss of equilibrium. 
The majority of the fish affected were striped mojarra (Diapterus 
plumieri). The tide at this time was low (ebb), and the water was 
clear. Along the high tide mark, D. plumeiri were frozen solid, an 
indication that the kill had begun as the tide receded. The re- 
maining fish collected dead at Bass Lake (table 1) included: brown 
bullhead (Ictalurns nebulosus), spotfin mojarra (Eucinostomus ar- 
genteus), fantail mullet (Mugil trichodon), and snook (Centropomus 
undecimalis). A seine-haul produced the following species which 
were unharmed: longnose killifish (Fundulus similis), gulf killifish 
(F. grandis), sheepshead minnow (Cyprinodon variegatus), rain- 
water killifish (Lucania parva), and blackcheek tonguefish (Syrnph- 
urus plagiusa). Juvenile red drum (Sciaenops ocellata), common 
in the area during November, were not collected or seen during 
the freeze. 

In Boca Ciega Bay the first fish kill was noted at 9 a.m., De- 
cember 14 (table 1). The surface water temperature was 10.4 °C 
and the salinity 32.97 o/oo. Crevalle jack (Caranx hippos) ap- 
peared to be affected in greater numbers than any other species in 
that area. We estimated that 300-500 jacks were killed in the 
areas sampled. These were found dead on the beaches and were 
netted, in a stunned condition, from the surface of the water. 
Those on the beaches appeared to have died as the tide receded. 
The fish collected ranged in size from 120 to 347 mm. but the 
majority were in two distinct size groups (120-150 mm. and 260- 
347 mm.). All specimens examined were immature. The second 
most numerous species affected in the same area was the lookdown 
(Selene vomer). Fourteen of the 17 specimens collected were 
dead on the beaches, while the others were taken from the sur- 
face of the water in a stunned condition. Additional species, col- 
lected dead from the beaches in Boca Ciega Bay, included 
lane snapper (Lutjanus synagris), striped burrfish (Chilomycterus 
schoepfi), planehead filefish (Monocanthus hispidus), cowfish (Lac- 
tophrys quadricornis), scaled sardine (Harengula pensacolae), spot- 
ted spoon-nose eel (Mystriophis intertinctus), and porcupinefish 
(Diodon sp.). 

Limited kills were observed at Point Pinellas and Big Bayou 
in central Tampa Bay on December 13 (table 1). Water tempera- 
ture was 11.2° C, and 10.8 °C, respectively, in the two areas. A 
seine-haul at Point Pinellas produced F. similis, F. grandis, C. 



14 Quarterly Journal of the Florida Academy of Sciences 

variegatus, tidewater silverside (Menidia beryllina), gulf pipefish 
(Syngnathus scovelli), and redfin needlefish (Strongylura notata), 
all in good condition. Young fantail mullet (M. trichodon), abun- 
dant at Pt. Pinellas during November, were not caught or observed 
during the freeze. 

On December 14, at Coffee Pot Bayou in central Tampa Bay 
(fig. 1), the water temperature was 11.8 °C at 5 p.m. The major 
species affected was C. hippos. Other species killed were permit 
(Trachinotus falcatus), Atlantic spadefish (Chaetodipterus faber), 
Harengula sp., and C. undecimalis (table 1). All dead fish were 
observed on the surface or near the shore. 

Discussion 

Fish kills associated with abnormal temperature reduction are 
common along the Florida Gulf coast (Storey and Gudger, 1936). 
The freeze of December 12-14 in the Tampa Bay area was the 
first and most severe cold spell of the winter. Air and water tem- 
peratures dropped rapidly. Nineteen fish species were adversely 
affected. The dominant fish killed were crevalle jack and striped 
mojarra. Cyprinodontids were tolerant of the low water tempera- 
tures. By December 17 water temperatures had increased in the 
sampling areas and no additional fish kills were observed. 

Springer and Woodburn (1960) reported minor fish kills in the 
Tampa Bay area on December 13, 1957, when water temperature 
was as low as 13 °C. Snook, barracuda, silver mullet, and striped 
mojarra were the predominant species affected. These fish kills 
occurred during another record cold winter when a low water 
temperature of 8.5 °C was reported (Dragovich, Finucane, and 
May, 1961). 

Storey (1937) observed that fish damaged most by a freeze at 
Sanibel Island, Florida, were tropical and subtropical species. 
Many of these species were also noted by the authors. They in- 
cluded: C. hippos, L. synagris, L. quadricornis, C. schoepfi, C. un- 
decimalis, M. intertinctus, and inshore lizardfish (Synodus foetens). 
Two species found by the authors but not included in Storey's dis- 
cussion were D. plumieri and S. vomer. Briggs (1960) gives the 
range of D. plumieri (Eugenes plumieri) as southwestern Florida 
to Bahia, Brazil, and west to Mexico. Springer and Woodburn 
(1960) stated that their collection of D. plumieri occurred in an 



Rinckey and Saloman: Cold and Marine Fishes 15 

area about 75 miles north of its reported range. This suggests 
that D. plumieri has a tropical and subtropical distribution. Joseph 
and Yerger (1956) listed S. vomer as being characteristic of trop- 
ical waters. The lookdown was not known to be common in Tampa 
Bay. Prior to the kill only three juveniles (28-43 mm.) were col- 
lected there in 18 months of sampling. All 17 specimens taken 
during the freeze were adults. 

Our observations suggest that the rapid, short term reduction 
in water temperatures of Tampa Bay had detrimental effects pri- 
marily on tropical and subtropical fish species. Other fish, in- 
cluding the majority of important sport and commercial species, 
were apparently unharmed. The total effect on local fish popula- 
tions appeared to be minor. 

Acknowledgment 

We wish to thank Mr. Martin Moe of the Florida State Board 
of Conservation, St. Petersburg, for data on the fish kill at Coffee 
Pot Bayou in central Tampa Bay. 

Summary 

Freeze conditions in central Florida during the period Decem- 
ber 12 through 14, 1962, lowered water temperatures rapidly in 
Tampa Bay. The effects of these conditions on Tampa Bay fish 
were noted. Nineteen fish species, the majority having tropical 
or subtropical distribution, were observed dead or stunned. Di- 
apterus plumieri and Caranx hippos suffered the greatest mortality. 

Literature Cited 

Briggs, John C. 1958. A list of Florida fishes and their distribution. Bull. 
Florida State Mus., Biol. Sci., vol. 2, no. 8, pp. 223-318. 

Dragovich, Alexander, John H. Finucane, and Billdz Z. May. 1961. 
Counts of red tide organisms, Gymnodinium hreve and associated 
oceanographic data from Florida west coast, 1957-59. U. S. Fish and 
Wildlife Serv., Spec. Sci. Rept., Fish. no. 369, pp. 1-175. 

Gunter, Gordon. 1941. Death of fishes due to cold on the Texas coast, 
January 1940. Ecology, vol. 22, no. 2, pp. 203-208. 

Gunter, Gordon, and Henry H. Hildebrand. 1951. Destruction of fishes 
and other organisms on the South Texas coast by the cold wave of 
January 28-February 3, 1951. Ecology, vol. 32, no. 4, pp. 731-736. 



16 Quarterly Journal of the Florida Academy of Sciences 

Joseph, Edwin B., and Ralph W. Yerger. 1956. The fishes of Alligator 
Harbor, Florida with notes on their natural history. Florida State 
Univ. Stud., no. 22, pp. 111-156. 

Springer, Victor G., and Kenneth D. Woodrurn. 1960. An ecological 
study of the fishes of the Tampa Bay area. Florida State Board 
Conserv., Prof. Pap. Ser., no. 1, pp. 1-104. 

Storey, Margaret. 1937. The relation between normal range and mor- 
tality of fishes due to cold at Sanibel Island, Florida. Ecology, vol. 
18, no. 1, pp. 10-26. 

Storey, Margaret, and E. W. Gudger. 1936. Mortality of fishes due to 
cold at Sanibel Island, Florida 1886-1936. Ecology, vol. 17, no. 4, 
pp. 640-649. 

Bureau of Commercial Fisheries Biological Station, St. Peters- 
burg Beach, Florida. Contribution No. 8. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



WESTERN ATLANTIC SERRANID FISHES (GROUPERS) 
OF THE GENUS EPINEPHELUS 

Luis Rene Rivas 

During the years 1945-47, the present writer, under a Guggen- 
heim Fellowship, spent several months studying the collections 
of western Atlantic groupers at the United States National Museum 
and the Museum of Comparative Zoology. Particular attention 
was paid to the genus Epinephelus, and extensive taxonomic in- 
formation was obtained. Numerous specimens representing all 
the presently recognized species of that group were examined. 
Since that time, material of all the western Atlantic species of 
Epinephelus has accumulated in the collections of the Ichthyolog- 
ical Museum of the University of Miami. Additional information 
was thus obtained, and during the summer of 1962 I studied the 
collections of Epinephelus at Stanford University. During the 
past 25 years, I have studied and collected in the field all the 
western Atlantic species of the genus. 

These studies were conducted in connection with a proposed re- 
vision but for various reasons publication was delayed. In the 
mean time Smith (1961), published a synopsis of the western north 
Atlantic groupers. In addition to biological and other data, his 
publication includes a key to the species of Epinephelus and re- 
lated genera. Although Smith's work is a valuable contribution 
to the knowledge of western Atlantic groupers, his key, the only 
means for identification given, is not satisfactory. One species, 
Epinephelus drummondhayi, was overlooked, and some of the key 
characters are misinterpreted or misleading. Furthermore, little 
indication is given of the usually considerable ontogenetic varia- 
tion of many of these characters. Some of the life color features 
used by Smith as key characters are of no value in the identification 
of preserved material. 

This paper is intended only as a mean for the identification of 
the western Atlantic species of Epinephelus as restricted by Smith 
(1961). For positive identification, the key should be used in con- 
junction with the diagnosis and remarks provided under each spe- 
cies heading. The species diagnoses combine most of the char- 
acters used in the key. The same characters, in the same sequence, 
are given in each diagnosis. Since nearly complete synonymies 



18 Quarterly Journal of the Florida Academy of Sciences 

and combinations were given by Smith, these will not be repeated 
in this study. 

In the dorsal and anal fin, the last two elements are counted as 
a single ray split to the base. All pectoral elements are counted. 
All gill rakers on the first arch are counted, including rudiments; 
the count for the lower limb includes the gill raker at the angle. 
Scales are difficult to count accurately and their number contributes 
little or nothing towards identification. For this reason scale 
counts have been omitted in this study. Color characters are 
based on specimens preserved in ethyl alcohol. Other characters 
are self-explanatory. Lengths are expressed in millimeters and 
refer to the standard length. 

I am grateful to Leonard P. Schultz, United States National 
Museum, William C. Schroeder, Museum of Comparative Zoology, 
and George S. Myers, Standard University, for permission to study 
and report on the material under their care. 

Genus Epinephelus Bloch 

Epinephelus Bloch, 1793, p. 11 (original description). Type species, E. mar- 
ginalis Bloch, 1793. 

The following combination of characters distinguishes the west- 
ern Atlantic species of Epinephelus from related genera occurring 
in the same area. 

Dorsal spines 10 or 11. Dorsal rays 13 to 18. Anal spines 3. 
Anal rays 8 to 10. Pectoral rays 16 to 19. Gill rakers on first arch 
13 to 19 (lower limb), 8 to 11 (upper limb), 21 to 28 (total). Supra- 
maxillary bone present and well developed. Preopercle without 
a strong antrorse spine at its angle. Pectoral fin not reaching to 
vertical from origin of anal fin. Caudal fin truncate, rounded or 
emarginate, not deeply forked. 

Key to Western Atlantic Species of Epinephelus 

la. Dorsal rays 13 to 15. Insertion of pelvic fin under or in advance of upper 

end of pectoral base; conspicuously in advance of lower end of 

pectoral base. Pelvic fin equal to or longer than pectoral, except 

in specimens about 400 mm. in length or larger. 

2a. Dorsal spines 10. Gill rakers 15 or 16, usually 15 on lower limb 

of first arch. Orbit diameter less than least interorbital width, 

except in very young specimens about 100 mm. in length or 



Rivas: Western Atlantic Groupers 19 

smaller. Caudal peduncle not conspicuously darker dorsally. 

1. Epinephelus nigritus 
2b. Dorsal spines 11. Gill rakers 15 to 17, usually 16 on lower limb of 
first arch. Orbit diameter equal to or greater than least inter- 
orbital width, except in specimens about 300 mm. in length or 
larger. Caudal peduncle conspicuously darker dorsally. 
3a. Dorsal rays 15. Posterior nostril 4 to 7 times larger than the 
anterior. Dusky band present around caudal peduncle but 
not forming a well-defined saddle-like black blotch dorsally. 
Dark-barred pattern on sides of body present. No white 
spots in regular rows on sides of body. 

2. Epinephelus mystacinus 

3b. Dorsal rays 13 to 15, usually 13 or 14. Posterior nostril about 

as large as the anterior or 3 to 5 times larger. Dusky band 

absent around caudal peduncle. A dorsal saddle-like black 

blotch on caudal peduncle gradually disappearing with age; 

faint or absent in specimens about 300 mm. in length or 

larger. Dark-barred pattern on sides of body absent. 

White spots in regular longitudinal and vertical rows on 

sides of body gradually disappearing with age; faint or 

absent in specimens about 300 mm. in length or larger. 

4a. Posterior nostril about as large as the anterior. Margin of 

spinous dorsal fin yellow. Dorsal saddle-like black 

blotch on caudal peduncle, when present (juveniles 

and young), not extending anteriorly to end of dorsal 

base or ventrally to lateral line. 

3. Epinephelus flavolimbatus 
4b. Posterior nostril 3 to 5 times larger than the anterior. Mar- 
gin of spinous dorsal fin dusky. Dorsal saddle-like 
black blotch on caudal peduncle, when present (juve- 
niles and young), extending anteriorly to end of dorsal 
base and ventrally to or beyond lateral line. 

4. Epinephelus niveatus 

ib. Dorsal rays 15 to 18, usually 16 or 17. Insertion of pelvic fin under or 

behind lower end of pectoral base; conspicuously behind upper end 

of pectoral base. Pelvic fin shorter than pectoral. 

5a. Gill rakers 13 to 15, usually 14 on lower limb of first arch. Orbit 

diameter less than least interorbital width, except in very young 

specimens about 150 mm. in length or smaller. Fifth dorsal 

spine less than least depth of caudal peduncle. Color pattern 

combining irregular dusky cross bars with brown spots. Size 

large; known to reach about 700 pounds in weight. 

5. Epinephelus itajara 

5b. Gill rakers 15 to 19, usually 16 to 18 on lower limb of first arch. 

Orbit diameter about equal to or greater than least interorbital 

width, except in specimens about 200 to 300 mm. in length or 



20 Quarterly Journal of the Florida Academy of Sciences 

larger. Fifth dorsal spine about equal to or greater than least 
depth of caudal peduncle. Color pattern not combining dusky 
cross bars with brown spots. Size medium to small; not known 
to reach 100 pounds in weight. 
6a. Anal rays 9 or 10, usually 9. Posterior margin of caudal fin 
straight or concave, except in young specimens about 150 
mm. in length or smaller. 
7a. Pectoral rays 18. Gill rakers 17 or 18 on lower limb of 
first arch. Dorsal fin membrane notched between the 
spines. No black specks scattered around eye. Head, 
body, and fins profusely speckled with white spots on 
a brown background. A dorsal saddle-like black blotch 
on caudal peduncle. 6. Epinephelus drummondhayi 
7b. Pectoral rays 16 to 18, usually 17. Gill rakers 15 or 16, 
usually 16 on lower limb of first arch. Dorsal fin 
membrane not notched between the spines. Black 
specks scattered around eye present, sometimes con- 
fined to preorbital and/or suborbital area. White 
spots, if present, on sides of body only; absent on head 
and fins. No dorsal saddle-like black blotch on cau- 
dal peduncle. 7. Epinephelus morio 
6b. Anal rays 7 to 9, usually 8. Posterior margin of caudal fin con- 
vex. 
8a. Gill rakers 16 to 19, usually 17 or 18 on lower limb of first 
arch. Head and body with numerous brown spots on 
a lighter background. Dark-barred pattern on sides of 
body absent. 
9a. Dorsal rays 16 or 17, usually 17. Pectoral rays 19, 
rarely 18. Spots on ventral area of body larger 
than those on back. A dorsal saddle-like black 
blotch on caudal peduncle. Three dark blotches 
along dorsal fin base. Soft dorsal, anal, and cau- 
dal fin not margined with dusky. 

8. Epinephelus adscensionis 
9b. Dorsal rays 15 or 16, usually 16. Pectoral rays 17, 
rarely 16. Spots on ventral area of body not 
larger than those on back. No dorsal saddle-like 
blotch on caudal peduncle. No dark blotches 
along dorsal fin base. Soft dorsal, anal, and cau- 
dal fin broadly margined with dusky. 

9. Epinephelus guttatus 
8b. Gill rakers 15 to 17, usually 16 on lower limb of first arch. 
Head and body without numerous brown spots on a 
lighter background; a few, widely scattered light spots 
sometimes present on sides of body. Dark-barred pat- 
tern on sides of body present or absent. 
10a. Dorsal rays 16 to 18, usually 17. Posterior nostril 



Rivas: Western Atlantic Groupers 21 

somewhat larger to about twice as large as the 
anterior. Black specks scattered around eye. 
Dark-barred pattern on sides of body present. A 
dorsal saddle-like black blotch on caudal ped- 
uncle. 10. Epinephelus striatus 
10b. Dorsal rays 15 or 16. Posterior nostril about as large 
as the anterior. No black specks scattered around 
eye. Dark-barred pattern on sides of body ab- 
sent. No dorsal saddle-like blotch on caudal ped- 
uncle. 11. Epinephelus guaza 

1. Epinephelus nigritus (Holbrook) 

Warsaw grouper; black grouper 

Serranus nigritus Holbrook, 1855, p. 173 (original description; Charleston, S. 
Carolina). 

Epinephelus nigritus, Smith, 1961, pp. 3 (characters in key), 11 (listed, syn- 
onymy in part; western north Atlantic). 

Diagnosis. Dorsal spines 10. Dorsal rays 13 to 15, usually 14. 
Anal rays 9. Pectoral rays 18 or 19, usually 18. Gill rakers on 
first arch 15 or 16, usually 15 (lower limb), 9 to 11, usually 9 or 10 
(upper limb), 24 to 26, usually 25 (total). Posterior nostril about 
as large as the anterior or somewhat larger. Orbit diameter less 
than least interorbital width; greater in very young specimens about 
100 mm. in length or smaller. Insertion of pelvic fin conspicuously 
in advance of upper end of pectoral base. Pelvic fin about equal 
to or longer than pectoral; shorter in specimens about 500 mm. in 
length or larger. Fifth dorsal spine about equal to or greater than 
least depth of caudal peduncle. Posterior margin of caudal fin 
convex. No black specks scattered around eye. Irregularly scat- 
tered white markings sometimes present on sides of body. Dark- 
barred pattern on sides of body absent. No dorsal saddle-like 
black blotch on nor dusky band around caudal peduncle. 

Remarks. Smith (1961, pp. 2, 3) and most authors before him 
have misinterpreted the number of dorsal spines in this species. 
Despite the statements "Dorsal spines usually 11 (sometimes 10 in 
Epinephelus nigritus)", or "Dorsal spines 10 or 11", the many speci- 
mens examined by me all had 10 dorsal spines. Perhaps Smith's 
misinterpretation was caused by his inclusion of Alphestes schol- 
anderi Walters (1957) in the synonymy of Epinephelus nigritus. 
As indicated by Walters in his original description and figures, the 



22 Quarterly Journal of the Florida Academy of Sciences 

posterior nostril is much larger than the anterior in scholanderi. 
Only Epinephelus mystacinus and E. niveatus have the posterior 
nostril enlarged. I have compared scholanderi with mystacinus 
and niveatus and have come to the conclusion that scholanderi is 
a synonym of niveatus, not of nigritus. The nostrils are about sub- 
equal in Epinephelus nigritus. 

2. Epinephelus mystacinus (Poey) 
Moustache grouper; cherna del alto 

Serranus mystacinus Poey, 1851, p. 52 (original description; Cuba). 
Epinephelus mystacinus, Smith, 1961, pp. 4 (characters in key), 11 (listed, 
synonymy; western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 15. Anal rays 9. 
Pectoral rays 18 or 19. Gill rakers on first arch 15 or 16, usually 
16 (lower limb), 8 or 9 (upper limb), 24 or 25 (total). Posterior 
nostril 4 to 7 times larger than the anterior. Orbit diameter about 
equal to or greater than least interorbital width; less in specimens 
about 400 mm. in length or larger. Insertion of pelvic fin under or 
slightly in advance of upper end of pectoral base. Pelvic fin about 
equal to or longer than pectoral; shorter in specimens about 400 
mm. in length or larger. Fifth dorsal spine greater than least depth 
of caudal peduncle; about equal to or somewhat less in specimens 
about 450 mm. in length or larger. Posterior margin of caudal 
fin convex. No black specks scattered around eye. No white 
spots in regular rows on body or fins. Dark-barred pattern on sides 
of body present. A dusky band around caudal peduncle much 
darker dorsally but not forming a well-defined saddle-like black 
blotch. 

Remarks. I do not agree with the name "misty grouper" given 
to this species by Smith (1961, p. 23). The scientific name mys- 
tacinus refers to the black, moustache-like band parallel to the 
upper jaw. For this reason, the name moustache grouper is con- 
sidered more appropriate for this species. 

Smith (1961, p. 4), in his key, attempts to distinguish mystacinus 
from niveatus and other species of Epinephelus on the basis of the 
enlarged posterior nostril. He overlooked, however, that in nive- 
atus the posterior nostril is also much enlarged and of the size 
stated by him for mystacinus. 



Rivas: Western Atlantic Groupers 23 

3. Epinephelus flavolimbatus Poey 

Yellowedge grouper; cherna del alto 

Epinephelus flavolimbatus Poey, 1865, p. 183 (original description; Cuba). 
Smith, 1961, pp. 3 (characters in key), 11 (listed, synonymy; western 
north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 13 to 15, rarely 15. 
Anal rays 9. Pectoral rays 18. Gill rakers on first arch 15 or 16, 
usually 16 (lower limb), 8 or 9 (upper limb), 24 or 25, usually 24 
(total). Posterior nostril about as large as the anterior. Orbit 
diameter about equal to or greater than least interorbital width; 
less in specimens about 450 mm. in length or larger. Insertion of 
pelvic fin under or in advance of upper end of pectoral base. Pelvic 
fin about equal to or longer than pectoral; shorter in specimens 
about 400 mm. in length or larger. Fifth dorsal spine about equal 
to or greater than least depth of caudal peduncle; less in specimens 
about 400 mm. in length or larger. Posterior margin of caudal fin 
convex in specimens up to about 300 mm. in length; straight or 
slightly concave in larger individuals. No black specks scattered 
around eye. Round white spots arranged in regular longitudinal 
and vertical rows on sides of body; sometimes a row on dorsal fin; 
these spots gradually disappear with age and are faint to absent 
in specimens about 300 mm. in length or larger. Dark-barred pat- 
tern on sides of body absent. A dorsal saddle-like black blotch on 
caudal peduncle not extending anteriorly to end of dorsal base or 
ventrally to lateral line; this blotch gradually disappears with age 
and is faint to absent in specimens about 300 mm. in length or 
larger. No dusky band around caudal peduncle. 

Remarks. This species resembles Epinephelus niveatus, with 
which it has been confused. Their color patterns are very similar, 
especially in preserved specimens up to about 300 mm. in length. 
Up to that size, however, the two species are well distinguished by 
the extent of the blotch on the back of the caudal peduncle. The 
coloration of the dorsal fin and the size of the nostrils distinguish 
the juveniles, young, and larger adults of flavolimbatus from those 
of niveatus. 



24 Quarterly Journal of the Florida Academy of Sciences 

4. Epinephelus niveatus (Valenciennes) 

Snowy grouper (young); golden grouper (adult); cherna del alto 

Serranus niveatus Valenciennes, in Cuvier and Valenciennes, 1828, p. 380 
(original description; Brazil). 

Epinephelus niveatus, Smith, 1961, pp. 3 (characters in key), 10 (listed, sy- 
onymy; western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 13 or 14, usually 14. 
Anal rays 9. Pectoral rays 18, rarely 19. Gill rakers on first arch 
15 to 17, usually 16 (lower limb), 8 to 10, usually 8 or 9 (upper limb), 
23 to 26, usually 24 or 25 (total). Posterior nostril 3 to 5 times 
larger than the anterior. Orbit diameter about equal to or greater 
than least interorbital width; less in specimens about 300 mm. in 
length or larger. Insertion of pelvic fin under or slightly in advance 
of upper end of pectoral base. Pelvic fin about equal to or longer 
than pectoral; shorter in specimens about 400 mm. in length or 
larger. Fifth dorsal spine greater than least depth of caudal ped- 
uncle; about equal to or somewhat less in specimens about 400 
mm. in length or larger. Posterior margin of caudal fin convex in 
specimens up to about 300 mm. in length; straight or slightly con- 
cave in larger individuals. No black specks scattered around eye. 
Round white spots arranged in regular longitudinal and vertical 
rows on sides of body; sometimes a row on dorsal fin; these spots 
gradually disappear with age and are faint to absent in specimens 
about 300 mm. in length or larger. Dark-barred pattern on sides 
of body absent. A dorsal saddle-like black blotch on caudal pedun- 
cle extending anteriorly to end of dorsal base and venrrally to or 
beyond lateral line; this blotch gradually disappears with age and 
is faint to absent in specimens about 300 mm. in length or larger. 
No dusky band around caudal peduncle. 

Remarks. As already indicated under Epinephelus nigritus, 
the form described as Alphestes scholanderi by Walters (1957) 
appears to be a synonym of Epinephelus niveatus not of E. nigritus. 
The close resemblance between niveatus and flavolimbatus was 
discussed under the latter species. 



Rivas: Western Atlantic Groupers 25 

5. Epinephelus itajara (Liechtenstein) 

Jewfish; spotted grouper; guasa 

Serranus itajara Liechtenstein, 1822, p. 278 (original description; Brazil). 

Epinephelus itajara, Smith, 1961, pp. 4 (characters in key), 12 (listed, synonymy; 
western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 15 or 16, usually 16. 
Anal rays 8. Pectoral rays 19. Gill rakers on first arch 13 to 15, 
usually 14 (lower limb), 8 or 9, usually 9 (upper limb), 21 to 24 
(total). Posterior nostril about equal to or somewhat larger than 
the anterior. Orbit diameter less than least interorbital width; 
greater in very young specimens about 150 mm. in length or small- 
er. Insertion of pelvic fin under or somewhat behind lower end 
of pectoral base. Pelvic fin shorter than pectoral. Fifth dorsal 
spine less than least depth of caudal peduncle. Posterior margin 
of caudal fin convex. No black specks scattered around eye; the 
markings on head are brown spots not restricted to area around 
eye. Head, body, and fins with dark-brown spots. Dark-barred 
pattern on sides of body present in specimens up to about 1000 
mm. in length. No dorsal saddle-like black blotch on caudal ped- 
uncle. A dusky band around caudal peduncle in young and half- 
grown. 

Remarks. The fewer gill rakers on the lower limb of the first 
arch and the color patern distinguish this species from all other 
western Atlantic members of the genus. Besides this species, 
only Epinephelus adscensionis and E. guttatus are brown-spotted 
but they have no cross bars. In E. itajara, the dark-barred pattern 
persists in specimens of a size seldom or never reached by adscen- 
sionis or guttatus. 

6. Epinephelus drummondhayi (Goode and Bean) 
Speckled hind; calico grouper 

Epinephelus drummondhayi Goode and Bean, 1878, p. 173 (original descrip- 
tion; Bermuda and Florida). Smith, 1961, p. 10 (listed; western north 
Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 15 or 16, usually 
16. Anal rays 9. Pectoral rays 18. Gill rakers on first arch 17 or 



26 Quarterly Journal of the Florida Academy of Sciences 

18 (lower limb), 9 or 10 (upper limb), 26 to 28, usually 27 (total). 
Posterior nostril about equal to or somewhat larger than the an- 
terior. Orbit diameter about equal to or greater than least inter- 
orbital width; less in specimens about 200 mm. in length or larger. 
Insertion of pelvic fin under lower end of pectoral base. Pelvic 
fin shorter than pectoral. Fifth dorsal spine about equal to or 
greater than least depth of caudal peduncle. Posterior margin of 
caudal fin straight or slightly concave. No black specks scattered 
around eye. Head, body, and fins profusely speckled with white 
spots on a brown background. Dark-barred pattern on sides of 
body absent. A dorsal saddle-like black blotch on caudal peduncle 
not extending anteriorly to end of dorsal base or ventrally to lat- 
eral line. No dusky band around caudal peduncle. 

Re?narks. This species was listed by Smith (1961, p. 10) but 
not included in his key. It is distinguished from all the other west- 
ern Atlantic species of the genus by the unique, striking color pat- 
tern. The truncate or emarginate caudal fin is another distinctive 
character shared only with Epinephelus morio. 

7. Epinephelus morio (Valenciennes) 

Red grouper; cherna americana; cherna de vivero 

Serranus morio Valenciennes, in Cuvier and Valenciennes, 1828, p. 285 (orig- 
inal description; New York; Santo Domingo). 

Epinephelus morio, Smith, 1961, pp. 2 (characters in key), 9 (listed, synonymy; 
western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 16 or 17. Anal 
rays 9, rarely 10. Pectoral rays 16 to 18, usually 17. Gill rakers 
on first arch 15 or 16, usually 16 (lower limb), 8 or 9 (upper limb), 
23 to 25 (total). Posterior nostril about equal to or somewhat larger 
than the anterior. Orbit diameter about equal to or greater than 
least interorbital width; less in specimens about 275 mm. in length 
or larger. Insertion of pelvic fin slightly to conspicuously behind 
lower end of pectoral base. Pelvic fin shorter than pectoral. Fifth 
dorsal spine about equal to or greater than least depth of caudal 
peduncle. Posterior margin of caudal fin convex in young up to 
about 150 mm. in length; straight or concave in larger individuals. 
Black specks scattered around eye, sometimes confined to preor- 
bital, suborbital area, or both. White spots sometimes present on 



Rivas: Western Atlantic Groupers 27 

sides of body. Dark-barred pattern on sides of body faint or 
absent. No dorsal saddle-like black blotch on, or dusky band 
around caudal peduncle. 

Remarks. This is the only western Atlantic species of Epineph- 
elus in which the dorsal fin membrane is not notched between the 
spines. In addition, it is distinguished from the other species, ex- 
cept E. drummondhayi, by the truncate or emarginate caudal fin. 
The fewer pectoral rays and gill rakers, and the color pattern dis- 
tinguish morio from drummondhayi. 

8. Epinephelus adscensionis (Osbeck) 

Rock hind; aguaji 

Trachinus adscensionis Osbeck, 1771, p. 96 (original description; Ascension 
Island). 

Epinephelus adscensionis, Smith, 1961, pp. 4 (characters in key), 11 (listed, 
synonymy; western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 16 or 17, usually 17. 
Anal rays 8. Pectoral rays 19, rarely 18. Gill rakers on first arch 
16 to 19, usually 17 or 18 (lower limb), 7 to 9 (upper limb), 23 to 
28, usually 25 to 27 (total). Posterior nostril about equal to the an- 
terior. Orbit diameter about equal to or greater than least inter- 
orbital width; less in specimens about 300 mm. in length or larger. 
Insertion of pelvic fin behind lower end of pectoral base. Pelvic fin 
shorter than pectoral. Fifth dorsal spine about equal to or greater 
than least depth of caudal peduncle. Posterior margin of caudal 
fin convex. No black specks scattered around eye; the markings 
on head are brown spots not restricted to area around eye. Head, 
body, and fins with brown spots which are larger on ventral area. 
Three dark blotches along dorsal fin base. Dark-barred pattern 
on sides of body absent. A dorsal saddle-like black blotch on cau- 
dal peduncle, not extending anteriorly to end of dorsal base or 
ventrally to lateral line; blotch faint or obsolete in specimens over 
300 mm. in length. No dusky band around caudal peduncle. 

Remarks. Smith (1961: 4) and several authors before him have 
stated that in this species there are no scales on the exposed surface 
of the maxillary. This has frequently been used as a distinguishing 
character but I have examined specimens with a well-defined patch 
of scales on the exposed surface of the maxillary. 



28 Quarterly Journal of the Florida Academy of Sciences 

9. Epinephelus guttatus (Linnaeus) 
Red hind; cabrilla 

Perca guttata Linnaeus, 1758, p. 292 (original diagnosis, after Catesby; Amer- 
ica). 

Epinephelus guttatus, Smith, 1961, pp. 3 (characters in key), 9 (listed, syn- 
onymy; western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 15 or 16, usually 16. 
Anal rays 7 or 8, usually 8. Pectoral rays 17, rarely 16. Gill rakers 
on first arch 17, rarely 16 (lower limb), 8 or 9, usually 9 (upper 
limb), 24 to 26 (total). Posterior nostril somewhat larger to about 
twice as large as the anterior. Orbit diameter about equal to or 
greater than least interorbital width. Insertion of pelvic fin under 
or slightly behind lower end of pectoral base. Pelvic fin shorter 
than pectoral. Fifth dorsal spine about equal to or greater than 
least depth of caudal peduncle. Posterior margin of caudal fin 
convex. No black specks scattered around eye; the markings on 
head are brown spots not restricted to area around eye. Head and 
body, and sometimes fins, with brown spots which are occasionally 
faint or absent on ventral area. Dark-barred pattern on sides of 
body absent. No dorsal saddle-like black blotch on, nor dusky 
band around caudal peduncle. 

Remarks. The "red spots" used by Smith (1961, p. 3) as a dis- 
tinguishing character for this species and for Epinephelus adscen- 
sionis refer to life color. These spots turn brown after a relatively 
short period of preservation in alcohol or formalin. 

10. Epinephelus striatus (Bloch) 

Nassau grouper; cherna criolla 

Anthias striatus Bloch, 1792, p. 125 (original description, after figure by Plu- 
mier; Atlantic Ocean). 

Epinephelus striatus, Smith, 1961, pp. 3 (characters in key), 8 (listed, synony- 
my; western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 16 to 18, usually 17. 
Anal rays 8. Pectoral rays 18, rarely 17. Gill rakers on first arch 
16 or 17, usually 16 (lower limb), 8 or 9, usually 9 (upper limb), 24 
to 26, usually 25 (total). Posterior nostril somewhat larger to 
about twice as large as the anterior. Orbit diameter about equal 



Rivas: Western Atlantic Groupers 29 

to or greater than least interorbital width; less in specimens about 
200 mm. in length or larger. Insertion of pelvic fin slightly to con- 
spicuously behind lower end of pectoral base. Pelvic fin shorter 
than pectoral. Fifth dorsal spine about equal to or greater than 
least depth of caudal peduncle. Posterior margin of caudal fin 
convex. Black specks scattered around eye. Dark-barred pattern 
on sides of body. A dorsal saddle-like black blotch on caudal 
peduncle, not extending anteriorly to end of dorsal base or ven- 
trally to lateral line. A dusky band, usually split into two, around 
caudal peduncle. 

Remarks. The figure of a 45.6 mm. specimen published by 
Smith (1961, p. 24) represents a juvenile which does not show the 
fully developed fins, body shape, or color pattern. 

11. Epinephelus guaza (Linnaeus) 
Grouper; garropa; guasa 

Labrus guaza Linnaeus, 1758, p. 285 (original diagnosis; "Pelago"). 
Epinephelus guaza, Smith, 1961, pp. 3 (characters in key), 13 (listed, synony- 
my; western north Atlantic). 

Diagnosis. Dorsal spines 11. Dorsal rays 15 or 16. Anal rays 
8 or 9, usually 8. Pectoral rays 18 or 19. Gill rakers on first arch 
15 or 16, usually 16 (lower limb), 8 to 10 (upper limb), 23 to 26 
(total). Posterior nostril about as large as the anterior or somewhat 
larger. Orbit diameter about equal to or greater than least inter- 
orbital width; less in specimens about 250 mm. in length or larger. 
Insertion of pelvic fin under or slightly behind lower end of pec- 
toral base. Pelvic fin shorter than pectoral. Fifth dorsal spine 
about equal to or greater than least depth of caudal peduncle. 
Posterior margin of caudal fin convex. No black specks scattered 
around eye. Irregularly scattered white spots sometimes present 
on sides of body. Dark-barred pattern on sides of body absent. 
No dorsal saddle-like black blotch on, nor dusky band around cau- 
dal peduncle. 

Literature Cited 

Bloch, M. E. 1793. Naturgeschichte des auslandischen Fische. Berlin 
(1785-1795), 9 parts, atlas. 

Cuvier, G., and A. Valenciennes. 1828. Histoire naturelle des poissons, 
vol. 2, 371 pp. Paris. 



30 Quarterly Journal of the Florida Academy of Sciences 

Goode, G. B., and T. H. Bean. 1878. On a new serranoid fish, Epinephelus 

drummondhayi, from the Bermudas and Florida. Proc. U.S. Nat. Mus., 
vol. 1, pp. 173-175. 

Holbrook, J. E. 1855. Ichthyology of South Carolina. Charleston, 182 
pp., 27 pis. 

International Game Fish Association. 1961. World record marine fishes. 
Miami, Florida, 15 pp. (unpaged). 

Lichtenstein, M. H. C. 1822. Die Werke von Marcgrave und Piso iiber die 
Naturgeschichte Brasiliens, erlautert aus den wieder aufgefundenen 
Originalzeichnungen. Abhandl. Akad. Wiss. Berlin (1821), pp. 267- 
288. 

Linnaeus, C. 1758. Systema naturae . . . etc., ed. 10, vol. 1, 824 pp. Hol- 
miae. 

Osbeck, P. 1771. A voyage to China and the East Indies . . . etc. London, 
2 vols. 

Poey, F. 1851. Especies nuevas de serranos, genero de peces de la familia 
de los percoideos. Mem. Hist. Nat. Isla de Cuba, vol. 1, pp. 50-60. 

. 1865. Peces nuevos de la Isla de Cuba. Rep. Fisico-nat. Isla de 

Cuba, vol. 1, pp. 181-192. 

Smith, C. L. 1961. Synopsis of biological data on groupers (Epinephelus 
and allied genera) of the western north Atlantic. FAO Fish. Biol. 
Synop., no. 23, sect. 1, 30 pp., 15 figs. 

Walters, V. 1957. Alphestes scholanderi, a new sea bass from the West 
Indies. Copeia, no. 4, pp. 283-286, 4 figs. 

Ichthyological Laboratory and Museum, Department of Zo- 
ology, University of Miami, Coral Gables, Florida. Contribution 
No. 51. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



FISHES OF SOME SOUTH CAROLINA COASTAL 
PLAIN STREAMS 

William D. Anderson, Jr. 

Distribution of the freshwater fishes in the Ashepoo, Comba- 
hee, Broad, and New River drainages of southeastern South Caro- 
lina was studied from 1956 through 1960. These systems are in 
the coastal plain between Edisto and Savannah River drainages 
in parts of Allendale, Bamberg, Barnwell, Beaufort, Colleton, 
Hampton, and Jasper counties. Ashepoo and Combahee drain 
into St. Helena Sound, Broad drains into Port Royal Sound, and 
New into Tybee Roads. Drainage areas of the rivers in square 
miles are: Ashepoo, 385; Combahee, 1,356; Broad, 727; and New, 
284; for a total of 2,752 square miles or 1,761,280 acres (communi- 
cation from Walter M. Bell, Corps of Engineers, Charleston, South 
Carolina). 

The general course of these rivers is to the southeast. For most 
of their lengths they flow over deposits of Tertiary age, but near 
their mouths they pass over sands and clays of Pleistocene or Re- 
cent age. At the headwaters of the Combahee the topography 
is somewhat rolling, but becomes flat a few miles southeastward 
and remains so, for the most part, all the way to the coast. 

This region is of considerable interest biologically, as it has a 
rich flora and fauna and is largely unspoiled by man. Much of 
the land is in the possession of large land owners who have been 
good guardians of our natural resources. This furnishes an ex- 
cellent comparison with areas not so well managed. Productive 
farms are scattered throughout; and a great portion of the territory 
is forested, some quite densely, making the pulp and timber in- 
dustry of great importance. There is little domestic or industrial 
pollution. The generally level lands lend themselves easily to 
good soil conservation practices, and as a consequence there is 
little erosion and almost no silting of streams. 

Most of the streams sampled are typical of the types which 
have their origins and entire courses confined to the South Atlantic 
coastal plain. ' They are usually clear and vary in color from that 
of drinking water (at or near their headwaters) to that of strong tea. 
The color stems from dissolved organic material. At a few stations 
(mostly stagnant pools) the water was milky or muddy. Bottom 



32 



Quarterly Journal of the Florida Academy of Sciences 



types encountered include muck, sand, gravel, and organic debris. 
The bottoms in still areas are mostly muck, but in areas of flowing 
water the bottoms are generally composed of soft to firm sand. 
At a few localities the bottom contains considerable gravel. Dead 
trees, limbs, stumps, and leaves make up an integral part of the 
bottom of many streams. In most cases, the streams are sur- 
rounded by swamps, predominantly gum-cypress, but which pos- 
sess a great diversity of flora. Aquatic plants are abundant (see 
Baldwin, 1956, p. 5-12, for a listing of the types of aquatics com- 
mon to this area). 

Ten-foot minnow seines and 20-foot bag seines were used to 
obtain most specimens. A few collections were made by poison- 
ing with rotenone. Each locality was assigned a station number. 
Specimens were preserved in the field in 10 percent formalin and 
were later converted to 38 percent isopropyl alcohol. 




Fig. 1. Map of South Carolina with study area crosshatched. 



Anderson: Fishes of South Carolina Streams 33 

Stations 

Collections were made from May 30, 1956, to May 25, 1959, 
mostly during the summers of 1956 and 1957. 

Choice of streams seined was limited by ease of access, and by 
depth of water and width of stream as only 10- and 20-foot seines 
were used. 

The list of stations contains: (1) station number, name of county, 
locality; (2) drainage; (3) date of collection; and (4) when available, 
certain environmental data, such as color and clarity of the water, 
most prominent bottom conditions encountered, and temperature 
of the water in degrees Fahrenheit at approximately 1 inch below 
the surface, taken in the shade, where possible. 

This study was not primarily ecological in nature, so most en- 
vironmental observations were qualitative and therefore subject to 
errors in judgment. These data present a more complete descrip- 
tion of the stations than locality alone. 

The field study was over a considerably larger area than is 
herein considered. Collections were made in brackish creeks in 
addition to fresh water. I am concerned here only with the species 
occurring in fresh water and have listed only the 120 freshwater 
stations, retaining the original field station numbers used in the 
overall survey. The general location of stations is shown in 
Figs. 1-2. 

2. Colleton Co.; where U.S. 15 crosses a creek; 4.5 miles N of Walter- 
boro; 5.5 miles S of Canadys. Ashepoo. May 30, 1956. Dark but clear. 
Muck. 

3. Colleton Co.; creek in North Walterboro crossed by U. S. 15; 3.2 
miles N of Walterboro; 1.25 miles S of station 2. Ashepoo. May 30, 1956. 
Dark but clear. Muck. 

4. Colleton Co.; where S-15-88 crosses one fork of the Ashepoo R.; 1.4 
miles E of U.S. 17A; 4.5 miles S of Walterboro. Ashepoo. May 30, 1956. 
Dark but clear. Muck. 

8. Bamberg Co.; Clear Pond on unmarked dirt road just off secondary 
road S-5-59; 0.5 of a mile E of U.S. 601; 6.4 miles SSE of Bamberg. Comba- 
hee. June 16, 1956. Clear. Sand and muck. 

9. Bamberg Co.; where U.S. 601 crosses Lemon Cr.; 1.25 miles S of 
Bamberg. Combahee. June 16, 1956. Dark but clear. Sand and muck. 

10. Bamberg Co.; where U.S. 301 crosses Grapevine Cr., a tributary of 
Lemon Cr.; 1.3 miles SW of Bamberg. Combahee. June 16, 1956. Rocks 
and muck. 



84 



Quarterly Journal of the Florida Academy of Sciences 




Fig. 2. The area studied in southeastern South Carolina. Station loca- 
tions are indicated by large black dots. 



11. Bamberg Co.; where U.S. 301 crosses the Little Salkehatchie R.; 
5.4 miles SW of Bamberg. Combahee. June 16, 1956. Dark but clear. 
Sand and muck. 

12. Bamberg Co.; where S-5-75 crosses Colston Branch, a tributary of 
the Little Salkehatchie R.; 9.4 miles S of Bamberg. Combahee. June 16, 1956. 
Dark but clear. Sand and muck. 

13. Bamberg Co.; where S-5-21 crosses the Little Salkehatchie R.; 11.6 
miles SSE of Bamberg; 4.3 miles ENE of Ehrhardt. Combahee. June 25, 
1956. Dark but clear. Sand and muck. 



Anderson: Fishes of South Carolina Streams 35 

14. Bamberg Co.; where S-5-87 crosses Savannah Cr.; 1.5 miles SW of 
Ehrhardt. Combahee. June 25, 1956. Dark but clear. Sand and muck. 

15. Colleton Co.; where S.C. 641 crosses Savannah Branch; 5.2 miles 
W of Lodge. Combahee. June 25, 1956. Dark but clear. Sand and muck. 

16. Allendale Co. -Bamberg Co.; where S.C. 641 crosses the Big Salke- 
hatchie R.; 5.5 miles SW of Ehrhardt. Combahee. June 25, 1956. Dark but 
clear. Muck. 

17. Bamberg Co.; where secondary road S-5-84 crosses Lemon Cr.; 8.8 
miles SSE of Bamberg. Combahee. June 26, 1956. Stagnant. Sand and 
muck. 

18. Bamberg Co.; where S-5-84 crosses the Little Salkehatchie R.; 8.9 
miles S of Bamberg. Combahee. June 26, 1956. Dark but clear. Sand. 

19. Bamberg Co.; where S-5-19 crosses Colston Branch; 5.4 miles E 
of Olar. Combahee. June 26, 1956. Dark but clear. Sand. 

20. Bamberg Co.; on unmarked dirt road just off S.C. 64, where it 
crosses Birds Branch; 0.6 mile SE of Olar. Combahee. June 26, 1956. Dark 
but clear. Sand and muck. 

21. Bamberg Co.; where U.S. 321 crosses the Little Salkehatchie R.; 
1.7 miles NNE of Govan. Combahee. June 26, 1956. Dark but clear. Rocks, 
sand, and muck. 

22. Bamberg Co.; where S.C. 70 crosses the headwaters of the Little 
Salkehatchie R.; 2.25 miles W of Denmark. Combahee. June 26, 1956. 
Dark but clear. Sand and muck. 

23. Bamberg Co.-Barnwell Co.; where S.C. 64 crosses Georges Cr.; 1.75 
miles WNW of Olar. Combahee. June 26, 1956. Dark but clear. Sand 
and muck. 

24. Bamberg Co.-Colleton Co.; where S.C. 217 crosses the Little Salke- 
hatchie R.; 5.25 miles ESE of Ehrhardt. Combahee. June 27, 1956. Sand 
and muck. 

25. Colleton Co.; where S-15-27 crosses Willow Swamp, a tributary 
of the Little Salkehatchie R.; 4.7 miles SW of Williams. Combahee. June 
27, 1956. Stagnant. Sand, muck, and organic debris. 

26. Colleton Co.; where S.C. 63 crosses the Little Salkehatchie R. at 
the first of four bridges W of U.S. 21; 9.7 miles S of Williams. Combahee. 
June 27, 1956. Stagnant. Muck and organic debris. 

27. Colleton Co.; overflow from a pond near Combahee R. just off a 
dirt road 0.25 of a mile SSE of U.S. 17A; 1.0 mile ENE of Yemassee. Comba- 
hee. July 23, 1956. Milky. Muck. 

28. Beaufort Co.-Colleton Co.; where dirt road ends at the Combahee 
R., just off U.S. 17A; 0.7 mile SE of station 27. Combahee. July 23, 1956. 
Clear. Sand and muck. 

29. Colleton Co.; where U.S. 21 crosses Black Cr.; 4.0 miles NW of 
U.S. 17; 4.5 miles N of Yemassee. Combahee. July 23, 1956. Clear. Sand 
and muck. 



36 Quarterly Journal of the Florida Academy of Sciences 

30. Colleton Co.; where S.C. 63 crosses the Little Salkehatchie R. at 
second bridge W of U.S. 21; 0.1 mile from station 26. Combahee. July 23- 
24, 1956. Clear. Muck and organic debris. 

31. Colleton Co.; where U.S. 17A crosses the Ashepoo R.; 0.5 mile WSW 
of Walterboro. Ashepoo. July 24, 1956. Dark but clear. Muck and organic 
debris. 

32. Colleton Co.; where U.S. 17 crosses Tupelo Swamp; 3.5 miles E of 
Ashepoo. Ashepoo. July 24, 1956. Dark but clear. Muck. 

35. Hampton Co.; where S.C. 63 crosses a tributary of the Big Salke- 
hatchie R. at second of three bridges; 5.5 miles ENE of Varnville. Combahee. 
July 26, 1956. Dark but clear. Sand and organic debris. 

36. Hampton Co.; where U.S. 601 crosses the Coosawhatchie R.; 1.4 
miles WSW of Hampton. Broad. July 26, 1956. Dark but clear. Sand 
and muck. 

37. Allendale Co. -Hampton Co.; where U.S. 321 crosses the Coosaw- 
hatchie R.; 2.5 miles WSW of Brunson. Broad. August 13, 1956. Dark 
but clear. Sand, muck, and organic debris. 

38. Hampton Co.; where S-25-49 crosses Blood Hill Cr., a tributary of 
Coosawhatchie R.; 3.5 miles SW of Brunson. Broad. August 13, 1956. 
Muddy. Muck. 

39. Hampton Co.; where S.C. 363 crosses the Coosawhatchie R.; 2.25 
miles WSW of Hampton. Broad. August 13, 1956. Muddy in spots, dark 
but clear in spots. Muck. 

40. Hampton Co.; where S.C. 128 crosses the Coosawhatchie R.; 2.9 
miles SSE of Varnville. Broad. August 13, 1956. Muddy in spots, dark but 
clear in spots. Muck. 

42. Hampton Co.; an old mill pond and spillway below pond, crossed 
by S-25-17; 1.9 miles W of Yemassee. Broad. August 14, 1956. Dark but 
clear. Muck. 

43. Hampton Co. -Jasper Co.; where S-25-36(S-27-87) crosses the Coosaw- 
hatchie R.; 3.7 miles SW of Early Branch. Broad. August 14, 1956. Dark 
but clear. Sand and muck. 

44. Jasper Co.; where S.C. 128 crosses Cypress Cr., a tributary of the 
Coosawhatchie R.; 1.0 mile NNW of Grays. Broad. August 15, 1956. Dark 
but clear. Muck. 

45. Jasper Co.; where S-27-108 crosses Cypress Cr., a tributary of Coosaw- 
hatchie R.; 2.3 miles NE of Grays. Broad. August 15, 1956. Dark but 
clear. Sand and muck. 

51. Bamberg Co.; where U.S. 601 crosses the Little Salkehatchie R.; 
9.3 miles S of Bamberg. Combahee. September 29, 1956. Clear. Sand 
and muck. 

52. Colleton Co.; where S-15-41 crosses a tributary of the Ashepoo R.; 
5.75 miles SE of Walterboro. Ashepoo. May 2, 1957. Clear. Muck. 64°. 



Anderson: Fishes of South Carolina Streams 37 

54. Beaufort Co. -Colleton Co.; where dirt road ends at the Combahee 
R., just off U.S. 17A; 0.7 mile SE of station 27. Combahee. May 2, 1957. 
Clear. Sand and muck. 68°. 

55. Barnwell Co.; where S-6-84 crosses Little Salkehatchie Cr.; 1.7 miles 
SE of Blackville. Combahee. June 10, 1957. Dark but clear. Sand and 
muck. 64°. 

56. Barnwell Co.; spillway below dam at Barnwell State Park, just off 
S.C. 3; 0.5 mile S of Blackville. Combahee. June 10, 1957. Dark but clear. 
Rocks, sand, and muck. 74°. 

57. Barnwell Co.; where S-6-67 crosses Turkey Cr.; 1.8 miles W of 
Blackville. Combahee. June 10, 1957. Dark but clear. Muck. 67°. 

58. Barnwell Co.; where S-6-168 crosses Turkey Cr.; 1.1 miles N of Barn- 
well. Combahee. June 10, 1957. Dark but clear. Sand. 67°. 

59. Barnwell Co.; where S.C. 28 crosses Buck Cr.; 4.0 miles NW of 
Barnwell. Combahee. June 10, 1957. Dark but clear. Sand. 67°. 

60. Barnwell Co.; where S.C. 28 crosses the spillway of a pond on 
Rosemary Cr.; 5.75 miles NW of Barnwell. Combahee. June 10, 1957. Dark 
but clear. Sand. 71°. 

61. Barnwell Co.; where S-6-185 crosses Rosemary Cr.; 1.25 miles S 
of Williston. Combahee, June 11, 1957. Dark but clear. Sand. 68°. 

62. Barnwell Co.; where S-6-166 crosses the Big Salkehatchie R.; 3.2 
miles NW of Barnwell. Combahee. June 11, 1957. Dark but clear. Sand. 
68°. 

63. Barnwell Co.; where S-6-20 crosses the Big Salkehatchie R.; 0.25 
mile from western boundary of Barnwell. Combahee. June 11, 1957. Dark 
but clear. Sand and muck. 68°. 

64. Barnwell Co.; where S.C. 28-37 crosses the Big Salkehatchie R.; 0.5 
mile S of Barnwell. Combahee. June 11, 1957. Dark but clear. Sand and 
muck. 70°. 

65. Barnwell Co.; where S.C. 64 crosses Hagood's Mill Cr.; 1.25 miles 
SE of Barnwell. Combahee. June 11, 1957. Dark but clear. Sand and 
muck. 68°. 

66. Barnwell Co.; where S-6-14 crosses Hercules Cr.; 4.5 miles ESE of 
Barnwell. Combahee. June 11, 1957. Dark but clear. Sand. 70°. 

67. Allendale Co.; where U.S. 301 crosses Log Branch; 0.9 mile NE 
of Allendale. Combahee. June 12, 1957. Dark but clear. Sand and 
muck. 70°.' 

68. Allendale Co.; where U.S. 321 crosses Jackson Cr., just below Tuten's 
Mill Pond; 0.75 mile SSE of Sycamore. Combahee. June 12, 1957. Dark 
but clear. Sand. 78°. 

69. Allendale Co.; where S-3-19 crosses the Coosawhatchie R.; 4.0 miles 
SSE of Allendale. Broad. June 12, 1957. Dark but clear. Sand. 72°. 

70. Allendale Co.; where S-3-47 crosses the Coosawhatchie R.; 1.6 miles 
S of Allendale. Broad. June 12, 1957. Dark but clear. Sand and muck. 
72°. 



38 Quarterly Journal of the Florida Academy of Sciences 

71. Allendale Co.; where S-3-22 crosses the Coosawhatchie R.; 0.3 mile 
SSE of Allendale. Broad. June 12, 1957. Dark but clear. Sand and 
muck. 78°. 

73. Jasper Co.; where S.C. 336 crosses a tributary of the New R.; 2.0 
miles SW of Ridgeland. New. June 13, 1957. Dark but clear. Sand 
and clay. 77°. 

74. Barnwell Co.; where S.C. 70 crosses Toby Cr.; 1.75 miles NE of 
Barnwell. Combahee. July 8, 1957. Clear. Sand. 70°. 

75. Barnwell Co.; where an unmarked dirt road crosses Hercules Cr.; 
3.7 miles E of Barnwell; 0.7 mile W of S-6-14. Combahee. July 8, 1957. 
Clear. Gravel and sand. 74°. 

76. Barnwell Co.; where S.C. 300 crosses Hurricane Cr.; 2.4 miles SE 
of Barnwell. Combahee. July 8, 1957. Clear. Sand. 72°. 

77. Barnwell Co.; where S-6-57 crosses the Salkehatchie R.; 6.25 miles 
S of Hilda. Combahee. July 8, 1957. Dark but clear. Sand. 77°. 

78. Allendale Co.; where S-3-48 crosses a small tributary of the Salke- 
hatchie R.; 3.75 miles ENE of Sycamore. Combahee. July 9, 1957. Clear. 
Sand. 73°. 

79. Allendale Co.; where S-3-18 crosses Jackson Cr.; 2.8 miles NE of 
Fairfax. Combahee. July 9, 1957. Dark but clear. Sand, muck, and organic 
debris. 76°. 

80. Allendale Co.; where S-3-69 crosses Duck Cr.; 1.8 miles SW of Fair- 
fax. Broad. July 9, 1957. Dark but clear. Sand and muck. 84°. 

81. Allendale Co.; where S.C. 641 crosses Miller Cr.; 0.3 mile E of 
Sycamore. Combahee. July 9, 1957. Dark but clear. Sand and muck. 75°. 

82. Allendale Co. -Barnwell Co.; where S.C. 300 crosses Wells Branch; 
1.4 miles NW of Ulmers. Combahee. July 10, 1957. Dark but clear. Sand 
and muck. 74°. 

83. Allendale Co.; where S.C. 28 crosses Log Branch; 0.25 of a mile 
N of Allendale. Combahee. July 10, 1957. Dark but clear. Sand and 
muck. 80°. 

84. Hampton Co.; where S-25-41 crosses Black Cr.; 3.9 miles SW of 
Hampton. Broad. July 10, 1957. Dark but clear. Gravel, sand, and 
muck. 79°. 

85. Hampton Co.; where an unmarked dirt road W of U.S. 321 crosses 
Black Cr.; 1.4 miles SW of Luray. Broad. July 10, 1957. Dark but clear. 
Gravel, sand, muck, and organic debris. 79°. 

86. Hampton Co.; where S.C. 631 crosses a tributary of Black Cr., be- 
tween Estill and Lena; 0.4 mile E of Estill. Broad. July 11, 1957. Dark 
but clear. Muck. 79°. 

87. Hampton Co.; where U.S. 601 crosses Briar Cr.; 4.1 miles NE of 
Estill. Broad. July 11, 1957. Dark but clear. Gravel and sand. 80°. 

88. Hampton Co.; where an unmarked dirt road off S.C. 631 crosses 
Cypress Cr.; 2.9 miles NE of Furman. Broad. July 11, 1957. Dark but 
clear. Sand. 75°. 



Anderson: Fishes of South Carolina Streams 39 

89. Hampton Co.; where an unmarked dirt road crosses Cypress Cr.; 
4.5 miles ENE of Furman. Broad. July 11, 1957. Dark but clear. Sand. 
77°. 

90. Hampton Co.; where S-25-94 crosses John Pen Brook; 0.9 mile E 
of Furman. Broad. July 12, 1957. Dark but clear. Gravel, sand, muck, 
and organic debris. 78°. 

91. Hampton Co.; where S-25-17 crosses Camp Brook; 3.3 miles ESE 
of Cummings. Broad. July 12, 1957. Dark but clear. Sand. 80°. 

92. Hampton Co.; where S-25-17 crosses a tributary of the Coosaw- 
hatchie R.; 3.7 miles W of Early Branch. Broad. July 12, 1957. Muddy. 
Sand. 76°. 

94. Colleton Co.; where S.C. 64 crosses Jones Swamp; 2.25 miles WNW 
of Walterboro. Ashepoo. July 23, 1957. Dark but clear. Sand, muck, 
and organic debris. 75°. 

95. Colleton Co.; where S-. 15-49 crosses Jones Swamp; 3.75 miles NW 
of Walterboro. Ashepoo. July 23, 1957. Muddy in spots; dark but clear in 
spots. Sand and muck. 74°. 

96. Colleton Co.; where S-15-89 crosses Jones Swamp; 2.75 miles NE 
of Stokes. Ashepoo. July 23, 1957. Dark but clear. Sand and organic 
debris. 77°. 

97. Colleton Co.; where an unmarked dirt road just off S-15-50 crosses 
Jones Swamp; 4.8 miles NNE of Stokes. Ashepoo. July 23, 1957. Muddy. 
Sand. 89°. 

98. Colleton Co.; where S-15-116 crosses Ireland Cr.; 4.6 miles NNE 
of Walterboro. Ashepoo. July 23, 1957. Dark but clear. Sand, muck, and 
organic debris. 75°. 

99. Colleton Co.; where U.S. 17A crosses Fuller Swamp; 5.0 miles E 
of Walterboro. Ashepoo. July 23, 1957. Dark but clear. Sand and muck. 
79°. 

100. Colleton Co.; where S-15-45 crosses Chessey Cr.; 1.75 miles SSE 
of Round O. Ashepoo. July 23, 1957. Dark but clear. Sand and muck. 

83°. 

103. Colleton Co.; where S-15-41 crosses the Ashepoo R. (Ivanhoe Cr.); 

3.7 miles NW of Green Pond; 0.8 mile SW of Ritter. Ashepoo. July 24, 
1957. Dark but clear. Muck. 86°. 

104. Colleton Co.; where S.C. 64 crosses Chessey Cr.; 4.4 miles ENE 
of Ritter. Ashepoo. July 24, 1957. Dark but clear. Muck. 81°. 

105. Colleton Co.; where S.C. 64 crosses Horseshoe Cr.; 4.5 miles NNE 
of Ashepoo. Ashepoo. July 24, 1957. Dark but clear. Muck. 80°. 

106. Colleton Co.; where S-15-41 crosses a tributary of Calfpen Swamp; 

3.8 miles NNW of White Hall. Combahee. July 24, 1957. Dark but clear. 
Sand and muck. 

107. Colleton Co.; where S-15-28 crosses Black Cr.; 4.25 miles WNW 
of Hendersonville. Combahee. July 24, 1957. Dark but clear. Muck. 



40 Quarterly Journal of the Florida Academy of Sciences 

108. Colleton Co.; where U.S. 21 crosses Sandy Run Cr.; 7.75 miles 
WNW of Hendersonville. Combahee. July 25, 1957. Dark but clear. Sand. 

109. Colleton Co.; where S.C. 64 crosses the Little Salkehatchie R. 
at the third of three bridges WNW of Bells Crossroads; 3.6 miles W of Ruffin. 
Combahee. July 25, 1957. Dark but clear. Sand. 

112. Colleton Co.; where U.S. 21 crosses Indian Cr.; 4.7 miles E of 
Islandton. Combahee. July 29, 1957. Dark but clear. Sand and muck. 74°. 

113. Colleton Co.; where S-15-33 crosses Indian Cr.; 5.25 miles SSE 
of Ruffin. Combahee. July 29, 1957. Dark but clear. Sand. 80°. 

114. Colleton Co.; where S.C. 63 crosses Deep Cr.; 2.5 miles ESE of 
Islandton. Combahee. July 29, 1957. Dark but clear. Muck. 75°. 

115. Colleton Co.; where S.C. 63 crosses Ricepatch Cr. at Islandton. 
Combahee. July 29, 1957. Dark but clear. Sand, muck, and organic 
debris. 75°. 

116. Colleton Co.; where U.S. 21 crosses Buckhead Cr.; 1.25 miles N 
of Ruffin. Combahee. July 31, 1957. Dark but clear. Sand and muck. 74°. 

117. Colleton Co.; where S-15-74 crosses Buckhead Cr.; 3.6 miles NNE 
of Ruffin. Combahee. July 31, 1957. Dark but clear. Sand. 76°. 

118. Colleton Co.; where S-15-63 crosses Buckhead Cr.; 2.3 miles NW 
of Smoaks. Combahee. July 31, 1957. Dark but clear. Sand, muck, and 
organic debris. 76°. 

119. Bamberg Co. -Colleton Co.; where S.C. 217 crosses a tributary of 
the Little Salkehatchie R.; 0.5 mile W of Padgetts. Combahee. July 31, 
1957. Dark but clear. Sand and muck. 76°. 

120. Colleton Co.; where S.C. 641 crosses Willow Swamp; 1.75 miles 
S of Lodge. Combahee. July 31, 1957. Dark but clear. Muck and clay. 
76°. 

121. Colleton Co.; where S-15-18 crosses Fender Cr.; just W of the 
center of Lodge. Combahee. July 31, 1957. Dark but clear. Sand and 
muck. 76°. 

122. Colleton Co.; where S-15-18 crosses a tributary of Willow Swamp; 
1.5 miles SW of Lodge. Combahee. July 31, 1957. Dark but clear. Sand, 
muck, and organic debris. 77°. 

123. Colleton Co.; where S-15-65 crosses Fuller Swamp; 5.1 miles ENE 
of Walterboro. Ashepoo. July 31, 1957. Dark but clear. Sand and muck. 
78°. 

124. Colleton Co.; where U.S. 17A crosses Chessey Cr.; 1.5 miles WNW 
of Cottageville. Ashepoo. July 31, 1957. Dark but clear. Sand and muck. 
77°. 

125. Hampton Co.; where S-25-44 crosses Deep Branch; 3.7 miles NNE 
of Early Branch. Combahee. August 1, 1957. Dark but clear. Sand, muck, 
and organic debris. 77°. 

126. Hampton Co.; where S-25-13 crosses Whippy Swamp; 2.9 miles 
SSE of Miley. Combahee. August 1, 1957. Dark but clear. Sand and 
muck. 76°. 



Anderson: Fishes of South Carolina Streams 41 

127. Hampton Co.; where U.S. 601 crosses Whippy Swamp; 1.25 miles 
WSW of Miley. Combahee. August 1, 1957. Dark but clear. Sand and 
muck. 80°. 

128. Hampton Co.; where S-25-43 crosses Whippy Swamp; 4.3 miles 
WNW of Miley. Combahee. August 1, 1957. Dark but clear. Sand, muck, 
and organic debris. 75°. 

129. Jasper Co.; where S.C. 631 crosses Cypress Cr. ; 2.0 miles WNW of 
Grays. Broad. August 12, 1957. Dark but clear. Sand and muck. 76°. 

130. Jasper Co.; where S.C. 128 crosses a tributary of the Coosawhatchie 
R.; 0.8 mile N of Gillisonville. Broad. August 12, 1957. Dark but clear. 
Sand and muck. 76°. 

132. Jasper Co.; where S-27-115 crosses a tributary of the headwaters 
of the New R.; 4.4 miles NW of Ridgeland. New. August 13, 1957. Dark 
but clear. Sand. 76°. 

133. Jasper Co.; where S-27-115 crosses a tributary of the headwaters 
of the New R.; 3.7 miles WNW of Ridgeland. New. August 13, 1957. Dark 
but clear. Muck. 82°. 

134. Jasper Co.; where S-27-39 crosses the New R.; 1.8 miles WNW of 
Ridgeland. New. August 13, 1957. Dark but clear. Sand and muck. 78°. 

135. Jasper Co.; where S-27-29 crosses the New R.; 1.5 miles WSW of 
Ridgeland. New. August 13, 1957. Dark but clear. Sand and muck. 80°. 

136. Jasper Co.; where U.S. 17 crosses the New R.; 1.6 miles SW of 
Switzerland. New. August 13, 1957. Dark but clear. Sand and muck. 78°. 

138. Jasper Co.; where S.C. 128 crosses Beaver Dam Cr.; 1.3 miles 
SE of Grays. Broad. August 14, 1957. Dark but clear. Muck. 78°. 

139. Jasper Co.; where an unmarked dirt road just off U.S. 17 crosses 
Bees Cr.; 2.25 miles NNE of Ridgeland. Broad. August 14, 1957. Dark 
but clear. Muck. 84°. 

140. Jasper Co.; where U.S. 17 passes alongside a drainage ditch, a 
tributary of the New R.; 2.2-2.3 miles SW of Switzerland. New. August 14, 
1957. Dark but clear. Muck. 95°. 

160. Colleton Co.; where S- 15-41 crosses the Ashepoo R. (Ivanhoe Cr.); 
3.7 miles NW of Green Pond; 0.8 mile SW of Ritter. Ashepoo. May 25, 1959. 
Dark but clear. Muck. (Same locality as station 103). 

Species of Fish 

No previous extensive work on the fishes of this area has been 
done. In several collections, Freeman (1954) obtained 14 species 
from a pond and a stream, north branch of the Salkehatchie River, 
Barnwell County, in the Atomic Energy Commission's Savannah 
River Plant Area. 

I collected and identified approximately 13,850 specimens rep- 
resenting 9 orders, 17 families, 31 genera, and 53 species. A 



42 Quarterly Journal of the Florida Academy of Sciences 

number of specimens were too small or too immature to be readily 
relegated to any species. Included in this group were members of 
the following genera: Erimyzon, Lepomis, Enneacanthus, Elassoma, 
and Etheostoma (Hololepis). The bulk of the collections is in the 
Charleston Museum, Charleston, South Carolina; but some speci- 
mens have been placed in the U.S. National Museum, Washing- 
ton, D.C. 

The list of species contains: order; family; scientific name and 
usually at least one common name; list of records of the species 
from the study area, if it has been previously reported (occasion- 
ally, after the list of previous records, references are included to 
works giving especially useful information on the particular spe- 
cies); number of specimens collected; and list of the stations from 
which the species was obtained. 

Order Semionotiformes 

Family Lepisosteidae 

Lepisosteus osseus (Linnaeus). Longnose gar 

37 specimens, from stations 26, 37, 84, 86, 91, 99, 104, 105, 109, 112, 
116-118, 120-121, 126, 128, 134. 

Order Amiiformes 
Family Amiidae 
Amia calva Linnaeus. Bowfin; Mudfish 
6 specimens, from stations 21, 88, 100, 129, 136, 138. 

Order Clupeiformes 
Family Umbridae 
Umbra pygmaea (DeKay). Eastern mudminnow 
25 specimens, from stations 52 and 100. 

Family Esocidae 

Esox americanus Gmelin. Redfin pickerel 

Esox americanus Gmelin. Freeman, 1954, p. 133, 148, Salkehatchie River. 

698 specimens, from stations 2-4, 9-15, 17-26, 30, 32, 35-40, 43-45, 52, 

56-57, 59, 64, 69, 73-74, 76, 78-91, 94-98, 100, 103-104, 108-109, 112-127, 

130, 133-135, 138-139, 160. 



Anderson: Fishes of South Carolina Streams 43 

Esox niger LeSueur. Chain pickerel 

Esox niger LeSueur. Freeman, 1954, p. 134, 148, Salkehatchie River. 

142 specimens, from stations 11, 13, 17-18, 21, 24, 29, 35, 39-40, 43-45, 
54, 77, 91, 94-95, 98, 103, 105, 107-109, 112, 114, 116-118, 120-122, 125, 
129-130, 160. 

Order Cypriniformes 

Family Cyprinidae 

Notemigonus crysoleucas (Mitchill). Golden shiner 

Notemigonus crysoleucas hosci (Valenciennes). Fowler, 1945, p. 168-170, 
Ashepoo River. 

Notemigonus hosci (Valenciennes). Fowler. 1935, p. 10-11, New River (prob- 
ably). 
1,068 specimens, from stations 2, 9-12, 14-15, 19, 25-26, 30, 32, 38-40, 

43-45, 60, 71, 73, 80, 83-92, 95, 97-100, 103-106, 112, 114-117, 119, 122-127, 

129-130, 132-136, 138-140. 

Opsopoeodus emiliae Hay. Pugnose minnow 

Opsopoeodus emiliae Hay. Anderson, MS, Combahee River (specimens listed 
below). 
26 specimens, from stations 14, 17, 18, 24, 35, 67, 79, 81. 

Notropis stonei Fowler. Sailfin shiner 

Notropis stonei Fowler. Suttkus, 1951, p. 83-84, Combahee and Broad Rivers. 

R. D. Suttkus now considers N. stonei as a subspecies of N. hypselopterus 

(personal communication). 
Erogala formosa (Putnam). Fowler, 1935, p. 16, Combahee River. 

980 specimens, from stations 9, 11-24, 29, 35, 37, 55, 57-59, 61-63, 65-67, 
69-70, 74-79, 81-82, 84, 89, 109, 126, 128. 

Notropis cummingsae Myers. Dusky shiner 

Notropis c. cummingsae Myers. Hubbs and Raney, 1951, p. 6-7, Combahee 

River. 

1,125 specimens, from stations 9, 11-19, 21-24, 30, 35, 43, 54, 58-63, 
65-67, 75-77', 79, 81-82, 109, 126, 128. 

Notropis chalyhaeus (Cope). Ironcolor shiner 

Notropis chalyhaeus (Cope). Fowler, 1945, p. 30, Little Salkehatchie River. 
Hydrophlox chalyhaeus (Cope). Fowler, 1935, p. 14, Combahee River. 
Cope, 1869, p. 383. 

1,149 specimens, from stations 9, 10-19, 21-25, 30, 35-37, 39, 43-45, 51, 
57, 60, 64, 71, 75, 77, 79, 81, 84-85, 88, 109, 112, 119, 126-127. 



44 Quarterly Journal of the Florida Academy of Sciences 

Notropis petersoni Fowler. Coastal shiner 

219 specimens, from stations 9-19, 21-25, 30, 35, 37, 54, 57, 60, 62-63, 
67, 79, 81-82, 126, 128. 

Notropis maculatus (Hay). Taillight shiner 

59 specimens, from stations 12, 25, 30, 32, 35, 40, 45, 51, 84, 109, 112, 
117, 121, 126-128. 

Family Catostomidae 

Minytrema melanops (Rafinesque). Spotted sucker 

Minytrema melanops (Rafinesque). Fowler, 1935, p. 10, Combahee River. 
11 specimens, from stations 15, 30, 57, 74, 84. 

Erimyzon sp. 

154 specimens, from stations 25, 30, 40, 70-71, 73, 77, 80, 89, 94-96, 
103, 105, 108-109, 112, 115-116, 125-126, 129, 134. 

Erimyzon sucetta (Lacepede). Lake chubsucker 

Erimyzon sucetta (Lacepede). Fowler, 1935, p. 9, New (probably) and Com- 
bahee Rivers. 

Erimyzon s. sucetta (Lacepede). Fowler, 1945, p. 168, Great Swamp Creek, 
49 miles south of Charleston on U.S. 17 (Ashepoo, probably). Freeman, 
1954, p. 134, 148, Salkehatchie River. 
100 specimens, from stations 26, 61, 83, 85, 87-88, 100, 117, 120, 122, 

130, 132-133, 135-136, 140. 

Erimyzon oblongus (Mitchill). Creek chubsucker 

263 specimens, from stations 2, 9-11, 19, 23, 38, 55, 84-86, 98, 119, 121- 
123, 127. 

Family Ictaluridae 

Ictalurus catus (Linnaeus). White catfish 

1 specimen, from station 105. 

Ictalurus platycephalus (Girard). Flat bullhead 
1 specimen, from station 30. 

Ictalurus natalis (LeSueur). Yellow bullhead 

Ameiurus natalis (LeSueur). Fowler, 1935, p. 18, Combahee River. 

81 specimens, from stations 4, 27, 29, 38, 40, 43-44, 68, 80, 84, 86-88, 95, 
99, 103-104, 108, 112, 116-117, 124-125, 129-130, 139. 



Anderson: Fishes of South Carolina Streams 45 

Ictalurus nebulosus (LeSueur). Brown bullhead 
12 specimens, from stations 59 and 60. 

Noturus gyrinus (Mitchill). Tadpole madtom 

Schilbeodes mollis (Hermann). Freeman, 1954, p. 138, 148, Salkehatchie 
River. (S. mollis is considered a synonym of N. gyrinus, personal com- 
munication with Reeve M. Bailey.) 

Hubbs and Raney, 1944, p. 25-26. 

46 specimens, from stations 4, 21, 30, 38, 44, 57, 64, 66, 68, 105, 116. 

Noturus leptacanthus Jordan. Speckled madtom 
7 specimens, from stations 23, 58-59, 81. 

Order Ancuilliformes 
Family Anguillidae 
Anguilla rostrata (LeSueur). American eel 
Muraena bostonensis LeSueur. Fowler, 1945, p. 167, Combahee River. 

56 specimens, from stations 4, 13-14, 24, 51, 69-70, 81, 88, 99, 123- 
124, 126. 

Order Cyprinodontiformes 

Family Cyprinodontidae 

Fundulus chrysotus (Gunther). Golden topminnow 

Zygonectes chrysotus (Gunther). Fowler, 1935, p. 20-21, New River (Prob- 
ably). 

Brown, 1957, p. 70-73, p. 75. 

Miller, 1955, p. 10. 

28 specimens, from stations 103 and 160. 

Fundulus notti (Agassiz). Starhead topminnow 

Fundulus notti lineolatus (Agassiz). Brown, 1958, p. 480, (Colleton County; 
Ireland Creek, tributary Ashepoo River), (Bamberg County; Little Salke- 
hatchie River between Denmark and Hilda, Route 70), (Jasper County; 
Great Swamp, tributary New River). 

Fundulus dispar lineolatus (Agassiz). Freeman, 1954, p. 139, 148, Salkehatchie 
River. 

Brown, 1957, p. 70-73, p. 76. 

Miller, 1955, p. 10. 

82 specimens, from stations 9, 25, 27, 39-40, 42, 44-45, 75, 83, 86, 91, 

94-95, 97-98, 103, 107-108, 112-113, 116-117, 125, 127, 129, 133-135, 140. 



46 Quarterly Journal of the Florida Academy of Sciences 

Family Poeciliidae 
Qambusia affinis (Baird and Girard). Mosquitofish 

Gambusia holbrooki (Agassiz). Fowler, 1945, p. 173, Combahee River and 

Great Swamp Creek, 49 miles south of Charleston on U.S. 17 (probably 

Ashepoo drainage). 
Gambusia ajfinis holbrooki (Girard). Freeman, 1954, p. 139, 148, Salkehatchie 

River. 

3,517 specimens, from stations 3, 4, 8-10, 12-15, 17, 19, 21, 23-25, 27-32, 
35-40, 42-45, 51-52, 60, 64, 67-69, 71, 77, 80-81, 84-88, 90-91, 94-100, 103- 
106, 108-109, 112, 114, 116-117, 119-120, 122-124, 126-127, 129-130, 132- 
136, 138-140, 160. 

Mollienesia latipinna LeSueur. Sailfin molly 
3 specimens, from station 32. 

Heterandria formosa Agassiz. Least killifish 
27 specimens, from stations 32, 52, 103, 106, 136, 160. 

Family Amblyopsidae 
Chologaster cornuta Agassiz. Swampfish 
6 specimens, from stations 2, 30, 73, 160. 

Order Percopsiformes 

Family Aphredoderidae 

Aphredoderus sayanus (Gilliams). Pirate perch 

Aphredoderus sayanus (Gilliams). Freeman, 1954, p. 139, 148, Salkehatchie 

River. 

404 specimens, from stations 2, 9, 13-15, 17, 19, 21-25, 29-30, 32, 35-36, 
38-40, 43-45, 51-52, 57, 59, 61, 64, 69-71, 77-87, 89, 94-96, 100, 103-105, 
108-109, 112-114, 116-117, 119-122, 124-127, 129-130, 132, 134-136, 139, 160. 

Order Perciformes 

Family Centrarchidae 

Micropterus salmoides (Lacepede). Largemouth bass 

Micropterus s. salmoides (Lacepede). Freeman, 1954, p. 140, 148, Salke- 
hatchie River. 

Bailey and Hubbs, 1949, p. 18-20. 

Hubbs and Bailey, 1940, p. 37-39. 

122 specimens, from stations 8-9, 19, 27-28, 30, 38, 40, 42, 45, 56, 60, 68, 

71, 74, 77, 79-80, 84, 91, 103-104, 109, 112, 116-117, 119-120, 125-126, 128- 

129, 132-136, 160. 



Anderson: Fishes of South Carolina Streams 47 

Chaenobryttus gulosus (Cuvier). Warmouth 
Chaenobryttus gulosus (Cuvier). Fowler, 1935, p. 25, Combahee River. 
Chaenobryttus coronarius Bartram. Freeman, 1954, p. 140, 148, Salkehatchie 

River. 

124 specimens, from stations 2, 9, 11, 13, 19, 21, 24-25, 30, 32, 38-40, 
43-45, 51-52, 56, 59-60, 69, 75, 79, 86, 95, 100, 104-105, 112, 116, 121, 124- 
125, 133-136, 138, 160. 

Lepomis sp. 
14 specimens, from stations 29, 39, 68, 84, 88, 109, 132, 134. 

Lepomis punctatus (Valenciennes). Spotted sunfish 

34 specimens, from stations 12-15, 17, 22, 24, 32, 35-36, 51, 58, 69, 80-81, 
96, 99, 109. 

Lepomis gibbosus (Linnaeus). Pumpkinseed 
11 specimens, from stations 15, 25, 26, 32, 122. 

Lepomis microlophus (Giinther). Redear sunfish; Shellcracker 
1 specimen, from station 56. 

Lepomis auritus (Linnaeus). Redbreast 

173 specimens, from stations 2, 9 ; 14, 18-19, 22, 25, 27-28, 37-38, 42-43, 
55, 57, 79, 82-85, 89, 95, 97-98, 105, 109, 117, 119-120, 123, 126, 128, 130, 
133-134, 136, 140. 

Lepomis marginatus (Holbrook). Dollar sunfish; Redeye 

[?] Lepomis megalotis (Rafinesque). Fowler, 1935, p. 24, New (probably) and 
Combahee rivers. 

Lepomis marginatus (Holbrook). Freeman, 1954, p. 141, 148, Salkehatchie 

River. 

165 specimens, from stations 2-3, 10-12, 22-23, 25, 30, 36-40, 43, 55, 
59-61, 75, 80-82, 84, 86, 89, 96-98, 100, 112, 115, 117-118, 120-123, 126-127, 
133-135, 140. 

Lepomis macrochirus Rafinesque. Bluegill; Bream 

Lepomis macrochirus purpurescens Cope. Freeman, 1954, p. 142, 148, Salke- 
hatchie River. 
508 specimens, from stations 8-13, 15, 17, 19, 21-22, 24-27, 30, 39-40, 

42-43, 51, 54-56, 59-60, 67-69, 71, 74, 80, 82-83, 85, 91, 98, 109, 121-122, 

125-127, 129-130, 133, 136, 140. 

Enneacanthus sp. 
17 specimens, from stations 83, 86, 95, 104. 



48 Quarterly Journal of the Florida Academy of Sciences 

Enneacanthus obesus (Girard). Banded sunfish 

168 specimens, from stations 2, 22, 25-26, 31-32, 36-37, 39, 43-44, 69, 
87, 96, 98-100, 103, 105-106, 114-117, 123-124, 133, 138-139. 

Enneacanthus gloriosus (Holbrook). Bluespotted sunfish 

41 specimens, from stations 21, 29, 37, 45, 69, 91, 94, 108, 127, 134- 
136, 160. 

Enneacanthus chaetodon (Baird). Blackbanded sunfish 

2 specimens, from station 29. 

Pomoxis nigromaculatus (LeSueur). Black crappie 
10 specimens, from stations 40, 80, 105, 140. 

Acantharchus pomotis (Baird). Mud sunfish 

45 specimens, from stations 21, 26, 30, 38-40, 43, 52, 86-87, 90-91, 94, 99, 
104, 106, 124, 133, 160. 

Centrarchus macropterus (Lacepede). Flier 

Centrarchus macropterus (Lacepede). Freeman, 1954, p. 142, 148, Salke- 

hatchie River. 

1,724 specimens, from stations 2, 9-12, 14-15, 19, 26, 29, 32, 36-40, 42-44, 
52, 59-60, 69, 71, 73, 79-80, 82-92, 94-95, 97, 99-100, 103-106, 108, 112-117, 
119, 123-125, 127, 129-130, 132-136, 138-139. 

Elassoma sp. 
1 specimen, from station 13. 

Elassoma zonatum Jordan. Banded pygmy sunfish 

Elassoma zonatum Jordan. Freeman, 1954, p. 143, 148, Salkehatchie River. 
15 specimens, from stations 12, 29-30, 44, 69, 85, 87, 103-104, 106, 109, 
117. 

Elassoma evergladei Jordan. Everglades pygmy sunfish 

Bohlke, 1956, p. 4. 

3 specimens, from stations 21, 87, 139. 

Family Percidae 
Percina nigrofasciata (Agassiz). Blackbanded darter 
[?] Alvordius peltatus (Cope). Fowler, 1935, p. 22, Combahee River. 
[?] Hadropterus peltatus crassus (Jordan and Brayton). Fowler, 1945, p. 195, 

Combahee River. 
Percina n. nigrofasciata x Percina n. raneyi (Intergrades). Crawford, 1956, p. 
50, Combahee River system. 
7 specimens, from stations 15, 24, 58, 75, 79, 82. 



Anderson: Fishes of South Carolina Streams 49 

Etheostoma olmstedi Storer. 

Etheostoma olmstedi. Cole, 1958, p. 1156, Santee-Cooper system southward 
to the St. Johns River. 
71 specimens, from stations 9, 12, 14-15, 17-18, 21-24, 59, 62, 66-67, 79, 

81-82. 

Etheostoma fricksium Hildebrand. Savannah darter 

Etheostoma fricksium Hildebrand. Anderson, MS, Combahee River (the speci- 
mens listed below). 

Hildebrand, 1923, p. 7-8. 

11 specimens, from stations 15, 23-24, 59, 76, 81-82. 



Etheostoma (Hololepis) sp. 
1 specimen, from station 82. 



Etheostoma serriferum (Hubbs and Cannon). Sawcheek darter 

Hololepis serrifer Hubbs and Cannon. Hubbs and Cannon, 1935, p. 31-36, 
Combahee River. Fowler, 1945, p. 196, Combahee River. 

Collette, 1962, p. 125-133. 

7 specimens, from stations 30, 36, 105, 109, 116. 

Etheostoma fusiforme barratti (Holbrook). Scalyhead darter 

Boleichthys fusiformis (Girard). Fowler, 1935, p. 23, Combahee River. 
Hololepis barratti (Holbrook). Hubbs and Cannon, 1935, p. 54-62, Combahee 

River. Fowler, 1945, p. 195, Combahee River. 
Etheostoma barratti (Holbrook). Freeman, 1954, p. 144, 148, Salkehatchie 

River. 
Collette, 1962, p. 150-190. 

5 specimens, from stations 52, 69, 95. 

Family Ephippidae 
Chaetodipterus faber (Broussonet). Atlantic spadefish 

Chaetodipterus faber (Broussonet). Anderson, MS, Ashepoo River (the speci- 
men listed below). 

Hildebrand "and Cable, 1938, p. 534-543. 
1 specimen, from station 103. 

Family Atherinidae 
Labidesthes sicculus (Cope). Brook silverside 

237 specimens, from stations 9, 11-13, 17, 21-22, 28, 30, 42-43, 45, 54, 
67-68, 80, 91, 94, 105, 112, 126, 128-129, 133-134. 



50 Quarterly Journal of the Florida Academy of Sciences 

Order Pleuronectiformes 
Family Soleidae 

Trinectes maculatus (Bloch and Schneider). Hogchoker 

Hubbs, 1932, p. 19-21. 

1 specimen, from station 105. 

Station of Special Interest 

It is impossible to discuss all collecting localities at length, but 
one is worthy of comment. Station 32 (Tupelo Swamp, Ashepoo 
drainage, Colleton County, July 24, 1956) is a typical, small, black- 
water creek. On the date of collection the stream was not flowing 
and was V2 to SV2 feet deep and 10 to 75 feet wide. Literally 
hundreds of fish were heard and observed breaking the surface 
of the water beneath the highway bridge and in its vicinity, and 
many Gambusia were seen swimming at the surface. About six 
or eight seine hauls were made, mostly beneath the bridge. Col- 
lecting at this station did not yield a great number of species, but 
many specimens were taken, particularly Gambusia and Centrar- 
chus. Centrarchus macropterus was the species largely heard and 
observed breaking the surface of the water. The following list 
gives the species and numbers of specimens collected: Esox ameri- 
canus, 3; Notemigonus crysoleucas, 43; Notropis maculatus, 2; Gam- 
busia affinis, 2,020 ±; Mollienesia latipinna, 3; Heterandria formosa, 
5; Aphredoderus sayanus, 1; Chaenobryttus gulosus, 34; Lepomis 
punctatus, 1; Lepomis gibbosus, 4; Enneacanthus obesus, 4; and 
Centrarchus macropterus, 542. 

Discussion 

Twenty-five species were collected that have not been reported 
from the area. Many other species were found that have not been 
recorded from one or more of the drainages. Table 1 gives a com- 
pilation of new records. 

All freshwater fish reported in the literature for this area were 
collected in at least one of the drainages studied. Fundulus chry- 
sotus and Etheostoma fusiforme barratti have been recorded from 
drainages in the area other than those in which I found them 
(Fundulus chrysotus from New and Etheostoma fusiforme barratti 
from Combahee). 



Anderson: Fishes of South Carolina Streams 



51 



TABLE 1 

New records of fish 



Species 



Entire 
Area 



Ashe- Comba- 
poo hee 



Broad New 



Lepisosteus osseus 


X 


X 


X 


X 


X 


Amia calva 


X 


X 


X 


X 


X 


Umbra pygmaea 


X 


X 








Esox americanus 




X 




X 


X 


Esox niger 




X 




X 




Notemigonus crysoleucas 






X 


X 




Notropis cummingsae 








X 




Notropis chalybaeus 








X 




Notropis petersoni 


X 




X 


X 




Notropis maculatus 


X 


X 


X 


X 




Minytrema melanops 








X 




Erimyzon sucetta 








X 




Erimyzon oblongus 


X 


X 


X 


X 




Ictalurus catus 


X 


X 


X 






Ictalurus platycephalus 


X 




X 






Ictalurus natalis 




X 




X 




Ictalurus nebulosus 


X 




X 






Noturus gyrinus 




X 




X 




Noturus leptacanthus 


X 




X 






Anguilla rostrata 




X 




X 




Fundulus chrysotus 




X 








Fundulus notti 








X 




Gambusia affinis 








X 


X 


Mollienesia latipinna 


X 


X 








Heterandria formosa 


X 


X 


X 




X 


Chologaster cornuta 


X 


X 


X 




X 


Aphredoderus sayanus 




X 




X 


X 


Micropterus salmoides 




X 




X 


X 


Chaenobryttus gulosus 




X 




X 


X 


Lepomis punctatus 


X 


X 


X 


X 




Lepomis gibbosus 


X 


X 


X 






Lepomis microlophus 


X 




X 






Lepomis auritus 


X 


X 


X 


X 


X 


Lepomis marginatus 




X 




X 




Lepomis macrochirus 




X 




X 


X 


Enneacanthus obesus 


X 


X 


X 


X 


X 


Enneacanthus gloriosus 


X 


X 


X 


X 


X 


Enneacanthus chaetodon 


X 




X 






Pomoxis nigromaculatus 


X 


X 




X 


X 


Acantharchus pomotis 


X 


X 


X 


X 


X 


Centrarchus macropterus 




X 




X 


X 


Elassoma zonatum 




X 




X 




Elassoma evergladei 


X 




X 


X 




Etheostoma serriferum 




X 




X 




Etheostoma fusiforme barratti 




X 




X 




Labidesthes sicculus 


X 


X 


X 


X 


X 


Trinectes maculatus 


X 


X 









52 Quarterly Journal of the Florida Academy of Sciences 

With the exception of Notemigonus crysoleucas in the Ashepoo 
and New drainages and Notropis maculatus in the Ashepoo, there 
was a scarcity of cyprinids in these two river drainages. Two fac- 
tors may have contributed to this. Experience has shown that the 
greater the number of areas sampled from a given drainage the 
greater the number of species one can expect to capture until all 
species occurring in the drainage have been collected. There were 
only 19 stations in the Ashepoo drainage and 7 in the New drain- 
age, as compared with 68 in the Combahee drainage and 26 in 
the Broad drainage. From the smaller number of stations seined 
in the Ashepoo and New drainages it might be expected that the 
total number of species collected from these systems would be 
smaller than the total number from the Combahee and Broad 
drainages. There was, however, not nearly so marked a difference 
in the number of centrarchid species obtained from the various 
drainages. Probably a more valid reason for the dearth of cypri- 
nids in the Ashepoo and New is that these drainages apparently 
lack stream environments which are especially conducive to habi- 
tation by cyprinids. The headwaters of the Combahee and Broad 
drainages flow through areas with considerably more elevation 
differential than do the headwaters of the Ashepoo and New drain- 
ages and, as a consequence, the Combahee and Broad drainages 
(especially the Combahee) have many relatively fast, small creeks, 
which may be called "minnow" streams. 

One specimen of Trinectes maculatus was taken in fresh water 
in the Ashepoo drainage. Hildebrand and Cable (1938, p. 630) 
and Gunter (1957, p. 14) mention several records of T. maculatus 
occurring in fresh water. 

Acknowledgments 

Numerous people helped in many ways during the course of 
this work. Foremost among these were Dr. Harry W. Freeman, 
formerly of the Department of Biology, University of South Caro- 
lina, who directed the research; Mr. William W. Anderson and Mr. 
Jack W. Gehringer, U.S. Bureau of Commercial Fisheries Biologi- 
cal Laboratory, Brunswick, Georgia, who gave many helpful sug- 
gestions in the preparation of the manuscript. Dr. Leonard P. 
Schultz, Curator of Fishes, U.S. National Museum, kindly allowed 
me to examine specimens. Mr. Curtis L. Smoak, Mr. William P. 



Anderson: Fishes of South Carolina Streams 53 

Clare, and others provided field assistance. The writer's father, 
Mr. W. D. Anderson, provided most of the necessary financial 
assistance. 

Literature Cited 

Anderson, William D., Jr. [MS] New records of fishes for South Carolina 
and the occurrence of Chaetodipterus faber in fresh water. 

Bailey, R. M., and C. L. Hubbs. 1949. The black basses (Micropterus) of 
Florida with descriptions of a new species. Occ. Pap. Mus. Zool., Univ. 
Mich., no. 516, pp. 1-40. 

Baldwin, William P. 1956. Food supply key to attracting ducks. S. Caro- 
lina Wildl., vol. 3, no. 1, pp. 5-12. 

Bohlke, James. 1956. A new pygmy sunfish from southern Georgia. Notul. 
Nat., Acad. Nat. Sci. Philadelphia, no. 294, pp. 1-11. 

Brown, J. L. 1957. A key to the species and subspecies of the cyprinodont 
genus Fundulus in the United States and Canada east of the continental 
divide. Jour. Wash. Acad. Sci., vol. 47, no. 3, pp. 69-77. 

. 1958. Geographic variation in southeastern populations of the cy- 
prinodont fish Fundulus notti (Agassiz). Amer. Midland Nat., vol. 59, 
no. 2, pp. 477-488. 

Cole, C. F. 1958. The taxonomy of the percid fishes of the genus Ethe- 
ostoma, subgenus Boleosoma, of eastern United States. Dissert. Abstr., 
vol. 18, no. 3, pp. 1155-1156. 

Collette, Bruce B. 1962. The swamp darters of the subgenus Hololepis 
(Pisces, Percidae). Tulane Stud. Zool., vol. 9, no. 4, pp. 115-211. 

Cope, E. D. 1869. Synopsis of the Cyprinidae of Pennsylvania. Trans. 
Amer. Philos. Soc, vol. 13, pp. 353-410. 

Crawford, R. W. 1956. A study of the distribution and taxonomy of the 
percid fish Percina nigrofasciata (Agassiz). Tulane Stud. Zool., vol. 4, 
no. 1, pp. 3-55. 

Fowler, H. W. 1935. Notes on South Carolina fresh water fishes. Contr. 
Charleston Mus., no. 7, pp. 1-28. 

. 1945. A study of the fishes of the southern Piedmont and Coastal 

Plain. Monogr., Acad. Nat. Sci. Philadelphia, no. 7, pp. 1-408. 

Freeman, H. W. 1954. Fishes of the Savannah River Operations Area. 
Univ. S. Carolina. Publ. Biol., Ser. 3, vol. 1, no. 3, pp. 117-156. 

Gunter, Gordon. 1957. Predominance of the young among marine fishes 
found in fresh water. Copeia, 1957, no. 1, pp. 13-16. 



54 Quarterly Journal of the Florida Academy of Sciences 

Hildebrand, S. F. 1923. Annotated list of fishes collected in vicinity of Au- 
gusta, Georgia, with description of a new darter. Bull. U.S. Bur. Fish., 
vol. 39, pp. 1-8. 

Hildebrand, S. F., and L. E. Cable. 1938. Further notes on the develop- 
ment and life history of some teleosts at Beaufort, N. C. Bull. U.S. 
Bur. Fish., vol. 48, no. 24, pp. 505-642. 

Hubbs, C. L. 1932. The scientific name of the common sole of the Atlantic 
coast of the United States. Proc. Biol. Soc. Wash., vol. 45, pp. 19-22. 

Hubbs, C. L., and R. M. Bailey. 1940. A revision of the black basses (Mi- 
cropterus and Huro) with descriptions of four new forms. Misc. Publ. 
Mus. Zool., Univ. Mich., no. 48, pp. 7-51. 

Hubbs, C. L., and M. D. Cannon. 1935. The darters of the genera Holo- 
lepis and Villora. Misc. Publ. Mus. Zool., Univ. Mich., no. 30, pp. 7-80. 

Hubbs, C. L., and E. C. Raney. 1944. Systematic notes on North American 
siluroid fishes of the genus Schilbeodes. Occ. Pap. Mus. Zool., Univ. 
Mich., no. 487, pp. 1-36. 

. 1951. Status, subspecies, and variations of Notropis cummingsae, 

a cyprinid fish of the southeastern United States. Occ. Pap. Mus. 
Zool., Univ. Mich., no. 535, pp. 1-25. 

Miller, R. R. 1955. An annotated list of the American cyprinodontid fishes 
of the genus Fundulus, with the description of Fundulus persimilis from 
Yucatan. Occ. Pap. Mus. Zool., Univ. Mich., no. 568, pp. 1-25. 

Suttkus, R. D. 1951. A taxonomic study of five cyprinid fishes related to 
Notropis hypselopterus of the southeastern United States. Ph.D. Thesis, 
Cornell Univ., 267 pp. 

U. S. Bureau of Commercial Fisheries Biological Laboratory, 
Brunswick, Georgia. Contribution Number 70. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



A PLIOCENE TEAL FROM SOUTH DAKOTA 
Pierce Brodkorb 

The living green-winged teal, Nettion crecca (Linnaeus), is a 
species of Holarctic distribution, represented in North America and 
Eurasia by slightly differentiated geographic races. In the Upper 
Pleistocene the species is recorded from no less than 32 sites in 
Europe, Asia, and North America (Brodkorb, 1964). N. bunkeri 
Wetmore (1944), a larger and apparently ancestral species, replaces 
it during the time interval near the Pleistocene-Pliocene boundary 
at several localities in western North America. In the Lower Plio- 
cene the presence of a large species in Germany, N. eppelsheimense 
(Lambrecht, 1933), and a very small one in Kansas, N. ogallalae 
Brodkorb (1962), suggests that more than one phyletic line of 
Nettion occurred in the northern hemisphere. The genus first 
makes its appearance in the Upper Miocene of France with N. 
velox (Milne-Edwards, 1867). 

The suggestion of a multiplicity of phyletic lines within the 
genus during the Tertiary is strengthened by a specimen for- 
warded to me for study by Dr. Morton Green of the South Dakota 
School of Mines and Technology. In some respects this Lower 
Pliocene fossil exhibits more similarity to certain southern hem- 
isphere teals that it does to the northern species. It is described 
below. 

Nettion greeni, new species 

Holotype. Distal half of right humerus (fig. 1), South Dakota 
School of Mines and Technology no. 63576. From Lower Pliocene 
in lower part of Ash Hollow Formation, at SDSM locality V631, 
D. C. Rice ranch, 3 miles northeast of Tu thill, Bennett County, 
South Dakota. Collected by Morton Green and Robert W. Wil- 
son, July 22, 1963. 

Diagnosis. Humerus agrees in characters with Nettion Kaup, 
as previously outlined (Brodkorb, 1962). Differs from living N. 
crecca (Linnaeus) and agrees with N. ogallalae from the Ogallala 
Formation of Kansas in having ectepicondyle rounded in lateral 
view and falling short of distal end of external condyle; largest 
foramen on palmer face close to tip of external condyle; scar of 
pronator longus with much medial thrust. 



56 Quarterly Journal of the Florida Academy of Sciences 



h \ 




Fig. 1. Nettion greeni, n. sp. Holotype humerus (actual length, 27.5 
mm.). 



Differs from N. ogallalae in having entepicondyle compressed 
in anconal view; proximal leg of entepicondyle straight in medial 
view (concave in N. ogallalae); pit for pronator bevis, in palmar 
view, in line with edge of shaft (bulging beyond shaft line in 
N. ogallalae), and in medial view pit long and mostly below level 
of upper end of facet for anterior articular ligament (pit rounded 
and mostly above level of upper end of facet in N. ogallalae): 
brachial depression very deep and extending distally nearly to 
facet for anterior articular ligament (in N. ogallalae, N. crecca, 
and the South American N. leucophrys brachial depression very 
shallow and distant from facet). 

Measurements. Distal width, 9.1; width of shaft, 4.3 mm. 
Size thus larger than the type of N. ogallalae, near minimum of 
N. crecca, and much smaller than N. eppelsheimense. The hu- 



Brodkorb: Pliocene Teal from South Dakota 57 

merus of N. bunker i and that of A 7 , velox are still unknown, but 
these are both relatively large species of teal. 

Discussion 

The functional significance of the enlargement, deepening, and 
distal encroachment of the brachial depression is unclear. M. 
brachialis, which originates here, flexes the forearm on the humerus 
and depresses the posterior edge of the wing. When contraction 
of M. triceps prevents flexion of the forearm, M. brachialis de- 
presses the posterior edge of the wing or holds the forearm hori- 
zontal against the great force of the air on the under surface of 
the secondaries (Fisher, 1946). Fisher was unable to correlate the 
development of M. brachialis with either soaring or flapping flight 
in vultures. Among the Pelecaniformes, however, I note that 
the brachial depression is large and deep and extends distally to 
the lower end of the facet for the anterior articular ligament in 
cormorants (Phalacrocorax), which flap, whereas it is shallow, 
shorter, and more proximal in darters (Anhinga), which soar. 

Some of the living southern hemisphere teals, notably Nettion 
flavirostre (Vieillot) and N. brasiliense (Gmelin) of South America 
and N. castanea (Eyton) of Australia, have a deep brachial depres- 
sion, although they differ from N. greeni in other characters. 

If an analogy to the Pelecaniformes could be made here in a 
different order, it would seem to suggest that N. greeni was a bird 
with even greater ability than other northern hemisphere teals to 
rise from the water in rapid, vertical flight. 

Acknowledgments 

I am indebted to Morton Green and Robert W. Wilson for 
the privilege of studying this specimen. Mr. and Mrs. D. C. Rice, 
of Tuthill, South Dakota, gave permission to the staff of the Mu- 
seum of Geology, South Dakota School of Mines and Technology, 
to camp and collect on their property. The photographs are by 
Robert W. McFarlane. The National Science Foundation spon- 
sored this paper through grant numbers G-19595 and GB-1686. 

Literature Cited 

Brodkorb, Pierce. 1962. A teal from the Lower Pliocene of Kansas. Quart. 
Jour. Florida Acad. Sci., vol. 25, no. 2, pp. 157-160, fig. 1. 



58 Quarterly Journal of the Florida Academy of Sciences 

. 1964. Catalogue of fossil birds. Part 2 (Anseriformes, Accipitri- 

formes, and Galliformes). Bull. Florida State Mus., in press. 

Fisher, Harvey Irvin. 1946. Adaptations and comparative anatomy of the 
locomotor apparatus of New World vultures. Amer. Midland Natural- 
ist, vol. 35, no. 3, pp. 545-727, figs. 1-28. pi. 1-13. 

Lambrecht, Kalman. 1933. Handbuch der Palaeornithologie. Gebriider 
Borntraeger, Berlin. 1024 pp., 209 figs. 

Milne-Edwards, Alphonse. 1867-1868. Recherches anatomiques et pale- 
ontologiques pour servir a l'histoire des oiseaux fossiles de la France. 
Victor Masson et fils, Paris. Vol. 1, 474 pp., 96 pi. 

Wetmore, Alexander. 1944. Remains of birds from the Rexroad fauna 
of the Upper Pliocene of Kansas. Univ. Kansas Sci. Bull., vol. 30, pp. 
89-105, figs. 1-19. 

Department of Biology, University of Florida, Gainesville, Flor- 
ida. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



TAPIRUS COPEI IN THE PLEISTOCENE OF FLORIDA 
Clayton E. Ray 

Remains of an unusually large tapir were found recently on the 
eastern shore of Tampa Bay directly across from St. Petersburg 
on the central Gulf coast of peninsular Florida. The material in- 
cludes the posterior portion of a left mandibular ramus with M 2 
well-preserved but with the roots only of M 3 and lacking the as- 
cending ramus, the posterior margin, and most of the horizontal 
ramus anterior to M 2 ; a right ramus with fragmentary symphyseal 
region, and lacking ascending and angular processes, but with 
P 2 -M 3 well-preserved (pi. 1); a few isolated fragments represent- 
ing incisors, a canine, and cheek teeth. The left and right rami 
are connected through a series of fragments which leave no doubt 
that they represent a single individual. It is probable that the 
remaining fragments also represent the same individual in that all 
were found in close proximity, and the size, preservation, and 
dental wear are similar throughout. 

The specimen was collected on August 31, 1963, by Lawrence 
A. Roberts and Kent M. Ainslie in a mound of dredged material 
in the extreme southwest corner of Sec. 22, T. 31 S., R. 19 E., 
Gibsonton Quadrangle, near Apollo Beach, Hillsborough County, 
Florida. The collectors very generously donated the material 
to the Florida State Museum, where it has been catalogued as 
UF 8225 (University of Florida Collections). In addition to the 
collectors, I wish to thank Dr. Horace G. Richards of the Academy 
of Natural Sciences of Philadelphia (ANSP) and Prof. Bryan Pat- 
terson of the Museum of Comparative Zoology (MCZ) for taking 
measurements of specimens in those institutions, and Dr. Malcolm 
C. McKenna of the American Museum of Natural History (AMNH) 
and Mr. Richard Ohmes of Chaires, Florida, for access to speci- 
mens in their care. 

Although the specimen was not found in situ, the source bed 
from which it was dredged is almost certainly the brown sand, 
equivalent to the Melbourne Formation, which has produced rich 
terrestrial vertebrate faunas from the Pinellas Peninsula (Seminole 
Field), Bradenton, and Sarasota. The light chocolate-brown stain- 
ing of the bone, darker brown staining of the dentine, and green 



60 Quarterly Journal of the Florida Academy of Sciences 




"'If 






■■■■■■■:::'::■:■:::■:.■■:'■:■'■.:::■'■;:■:■': ■■■-;■■-■,-•■:-■■—■■, ■■■; , ■■ ■:■■■: 








B 
























Plate 1. Right mandibular ramus of Tapirus copei. UF 8225, in lingual 
(A), occlusal (B), and labial (C) aspects. Approximately X 0.4. 



Ray: Cope's Tapir in the Florida Pleistocene 61 

to blue-black mottled staining of the enamel are characteristic 
of specimens from this horizon in the area. 

The specimen greatly exceeds in size all jaws of Tapirus avail- 
able to me from Florida (table 1), and is exceeded in size among 
North American Pleistocene tapirs probably only by the poorly 
known T. merriami Frick from southern California. The dimen- 
sions of individual teeth are compared (table 2) with samples ana- 
lyzed by Simpson (1945) of T. veroensis from the Seminole Field 
and T. copei from the Port Kennedy fissure in southeastern Penn- 
sylvania. UF 8225 lies within the observed range for the sample 
of T. veroensis in only 3 of 17 variates of the lower cheek teeth, 
P 3 WA, P 4 WP, P 4 WP, and within the standard range in one (MiL) of 
9 variates for which the standard range was calculated. Abbrevi- 
ations for tooth dimensions are those of Simpson. Application of 
Student's f-test to each of the 9 variates available for the lower 
molars, indicates that in every case the probability is essentially 
nil (P less than .025) that UF 8225 is drawn from a population 
with a mean no greater than that of the population represented by 
the Seminole Field sample. 

UF 8225 lies at or above the mean for the sample of T. copei 
in all but one of 17 variates (M 3 WA). It lies within the observed 
range for T. copei in all but 4 of the 17 variates (P 2 L, P 2 W, P 3 WP, 
M 3 L). Two of these (P 3 WP, M 3 L) fall within the standard range 
for T. copei, but both dimensions of P 2 lie above the standard 
range for T. copei. Testing the hypothesis that the mean of the 
population from which UF 8225 is drawn is not different from 
that of the population from which the sample of T. copei is drawn, 
shows (table 2) that deviations as great or greater than those ob- 
served might be expected in 5 per cent or more of similar trials 
for all of the 17 variates save 3 (P 2 L, P 2 W, P 3 WP). 

It is obvious from the above that UF 8225 is much closer to 
the sample of T. copei in dental dimensions than it is to the sample 
of T. veroensis, and in fact lies near or beyond the extreme of 
variation in T. copei farthest from T. veroensis. The biological 
significance of these facts and the proper taxonomic assignment of 
UF 8225 remain to be determined. 

Bader (1957, p. 70) has pointed out quite correctly that a "sta- 
tistical treatment of all teeth now available from Florida would 
probably increase the variation somewhat over that reported by 
Simpson." This procedure would be applicable only to faunas 



Quarterly Journal of the Florida Academy of Sciences 



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Ray: Cope's Tapir in the Florida Pleistocene 63 

(such as those from the Melbourne bone bed) for which there is 
reasonable assurance of contemporaneity. Bader (1957, p. 70) 
erroneously challenged the supposed specific separation of the 
rather small AMNH 2311, for Simpson (1945, p. 64) explicitly re- 
garded the specimen as conspecific with T. veroensis. Simpson 
(p. 64) based his statement, "It is . . . demonstrable that at least 
one other kind of tapir . . . occurred in Florida," not upon small, 
but upon large specimens, such as AMNH 23110 (M 3 L 30.5 WA 
21.5, WP 19.6), a mandibular fragment without exact locality re- 
ferred by Simpson tentatively to T. copei, but not discussed by 
Bader. Even if all other mandibles of Tapirus known from Florida 
were brigaded as a single population of T. veroensis, it is improb- 
able that the assuredly expanded variation would be sufficiently 
great to permit interpretation of AMNH 23110 and UF 8225 on 
the basis of individual variation. 

Sexual dimorphism would seem to be out of the question in 
this case, in view of the paucity of large, hypothetically male, 
individuals in Florida and apparent absence of small individuals 
at Port Kennedy, and the lack of evidence for sexual dimorphism 
in modern tapirs (Simpson, 1945, p. 42; Hooijer, 1947, p. 297). 

Geographic variation cannot be called upon to explain the 
differences between UF 8225 and the Seminole Field sample, for 
the localities are within 25 miles of one another across Tampa Bay. 
Even if there were a Pinellas Island at the time, as suggested by 
Cooke (1945, pp. 298, 308), its isolation is unlikely to have been 
sufficiently profound or prolonged to generate divergence in the 
tapir population. Further, Pleistocene tapirs from Florida as a 
whole are predominantly of the smaller, T. veroensis, size. 

Leaving UF 8225 aside for the moment, geographic variation 
could be involved in the difference in size between T. copei from 
Port Kennedy and T. veroensis from Seminole Field. The two 
samples might be regarded as vicariant species or as vicariant pop- 
ulations of a single species, with the larger form more northerly 
as might be expected. Although Simpson (1945, p. 42) found no 
evidence of geographic differentiation in modern T. terrestris over 
an area larger than the eastern United States, he did not have 
large local samples, and it may be also that Pleistocene tapirs lived 
in the United States under a much more radical latitudinal climatic 
gradient than does T. terrestris in South America today. How- 
ever, the two samples cannot more than suggest geographic varia- 



64 



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Ray: Cope's Tapir in the Florida Pleistocene 65 

tion as a factor, for they definitely are not coeval: Port Kennedy, 
Yarmouthian (Kurten, 1963, pp. 99, 101); Seminole Field, Wiscon- 
sinan (Weigel, 1962, pp. 52, 53). 

If, as seems at least probable, UF 8225 is derived from the 
same horizon as the Seminole Field fauna, and if it represents 
T. copei, then it provides evidence that T. copei and T. veroensis 
are not vicariant populations, but are specifically distinct and 
were sympatric in Florida, although T. veroensis must have been 
by far the more common species. The palate and isolated teeth 
recorded from Melbourne by Gazin (1950, p. 403) as Tapirus cf. 
haysii may provide the best test for sympatry of T. copei and T. 
veroensis in Florida, but this cannot be evaluated until the rich 
sample from Melbourne has been analyzed. 

Meanwhile the possibility must be considered that UF 8225 
is not coeval with the Seminole Field fauna or with any tapir 
(save perhaps AMNH 23110) previously recorded in Florida, and 
represents an interval (glacial?) rare, or rarely tapped, in the 
Floridian record, when T. copei was present. Under these cir- 
cumstances, T. copei and T. veroensis could be regarded as con- 
specific, perhaps representing a chronocline of decreasing size 
analogous to that indicated by Hooijer (1947, p. 290) leading from 
T. indicus intermedins to the living T. indicus indicus, but not 
including the larger, apparently more advanced, Megatapirus au- 
gustus (see Colbert and Hooijer, 1953, p. 90). The discrepancy in 
tooth dimensions between these two forms (see Hooijer, p. 291), 
regarded by Hooijer as subspecies, is at least as great as that be- 
tween T. copei and T. veroensis. Under this interpretation the 
differences between T. copei (including UF 8225 and AMNH 
23110) and T. veroensis would include both geographic and tem- 
poral components. Only on such a hypothesis is the suggestion 
(Bader, 1957, p. 70; Weigel, 1962, p. 41) supportable on present 
evidence that a single species of tapir occurs in the Pleistocene 
of Florida.. 

Knowledge of the cranial characters of T. copei and the dis- 
covery or analysis of additional known-age samples of Tapirus 
in eastern North America will aid in determining whether T. copei 
and T. veroensis are collateral species or successive stages in a 
chronocline. In any case, the specimen from Apollo Beach should 
be assigned to T. copei, whether the species is valid or is shown 
eventually to be a subspecies of T. veroensis. The fact that the 



66 Quarterly Journal of the Florida Academy of Sciences 

dimensions of P 2 fall above the standard range for the Port Ken- 
nedy sample in my opinion does not merit taxonomic recognition 
but serves to emphasize the present inadequacy of data on Pleisto- 
cene tapirs in North America and the hazards of assigning isolated 
teeth of tapirs to species. 

Literature Cited 

Bader, R. S. 1957. Two Pleistocene mammalian faunas from Alachua 
County, Florida. Bull. Florida State Mus.. vol. 2, no. 5, pp. 53-75, 
figs. 1-6. 

Colbert, E. H., and D. A. Hooijer. 1953. Pleistocene Mammals from the 
limestone fissures of Szechwan, China. Bull. Amer. Mus. Nat. Hist., 
vol. 102, art. 1, pp. 1-134, text-figs. 1-42, pis. 1-40. 

Cooke, C. W. 1945. Geology of Florida. Geol. Bull. Florida Geol. Surv., 
no. 29, ix + 339 pp., text-figs. 1-47, pi. 1. 

Gazin, C. L. 1950. Annotated list of fossil Mammalia associated with human 
remains at Melbourne, Fla. Jour. Washington Acad. Sci., vol. 40, 
no. 12, pp. 397-404. 

Hooijer, D. A. 1947. On fossil and prehistoric remains of Tapirus from 
Java, Sumatra and China. Zool. Mededeelingen, vol. 27, pp. 253- 
299, pis. 1-2. 

Kurten, B. 1963. Notes on some Pleistocene mammal migrations from the 
Palaearctic to the Nearctic. Eiszeitalter und Gegenwart, vol. 14, pp. 
96-103, figs. 1-2. 

Simpson, G. G. 1945. Notes on Pleistocene and Recent tapirs. Bull. Amer. 
Mus. Nat. Hist., vol. 86, art. 2, pp. 33-82, text-figs. 1-11, pis. 5-10. 

Weigel, R. D. 1962. Fossil vertebrates of Vero, Florida. Special Publ. 
Florida Geol. Surv., no. 10, vii + 59 pp., figs. 1-6. (First copies mailed 
January 22, 1963). 

Florida State Museum and Department of Biology, University 
of Florida, Gainesville, Florida (present address: U. S. National 
Museum, Washington, D. C). 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



NOTES ON THE ODONATA OF CUBA 

MlNTER J. WESTFALL, Jr. 

The island of Cuba has a very rich odonate fauna which to 
date has been studied inadequately. The eminent Cuban natural- 
ist, Juan Gundlach, made a noteworthy contribution to the knowl- 
edge of Cuban species when from 1888 to 1890 he included the 
Odonata in Part 3 of the Contribution a la Entomologia Cubana. 
There he listed 70 species. The paper was almost completely over- 
looked until Dr. Philip P. Calvert made it known through his criti- 
cal study in 1919. He reduced Gundlach's list to 69, including one 
subspecies. 

In 1931 Richard Dow published the results of several weeks' 
collecting during August and September of 1930, chiefly at Cen- 
tral Soledad, the sugar estate on which the Harvard Botanical 
Garden is located. This estate is near Cienfuegos, Las Villas 
Province (referred to as the Province of Santa Clara in Dow's 
paper). His list contained 35 species. 

Dr. James G. Needham spent the month of April, 1937 at the 
same location, alluding to it as the "Atkins Institution of Harvard 
University" in his paper of 1941. Accompanied by Dr. J. C. Brad- 
ley, Dr. Needham again visited Cuba during the last ten days of 
March, 1939, collecting at Central Soledad as well as in the 
Province of Pinar del Rio. As a result of this trip Dr. Needham 
(1939) described the nymph of Neoneura carnatica Selys. He told 
me that he went into the Trinidad Mountains hoping to get the rare 
Microneura caligata Selys but was unsuccessful in finding it on 
either trip to Cuba. I now find from examining his collections that 
he took some nymphs of this species in 1939 but did not recog- 
nize them. 

It was with this endemic and elusive damselfly in mind that 
my wife and I went to Cuba for two weeks late in May 1959 with 
the aid of a grant from the National Science Foundation. We also 
chose the Atkins Botanical Garden as headquarters, and when we 
arrived on May 27 were made very much at home in "Harvard 
House" by Dr. and Mrs. Duncan Clement. Until we left on June 
9, we spent busy days collecting and rearing Odonata. Much of 
the time was spent at the Garden collecting around ponds and 
pools of a variable stream. 



68 Quarterly Journal of the Florida Academy of Sciences 

By automobile we made trips to the nearby Trinidad Moun- 
tains, stopping at several streams and ponds along the way. Our 
total collection included 879 adult specimens, representing 44 spe- 
cies, and numerous nymphs and exuviae. Because several of 
these were omitted from Dow's list of Soledad, I am giving a 
complete list of all species taken. Reference is made under each 
to the localities listed below by number. Eleven species of Anisop- 
tera and nine of Zygoptera were reared. 

Two other collections of Cuban Odonata have been studied 
and are being included [within brackets to differentiate our 1959 
collection], making the total species reported in this paper 52. 
The United States National Museum (USNM) collection is made 
up of specimens collected by E. A. Schwarz in 1904 and Charles T. 
Ramsden in 1946, chiefly from Cayamas and Santiago de Cuba in 
Oriente Province. Dr. Calvert's collection from the Academy of 
Natural Sciences of Philadelphia (ANSP) contains specimens from 
two localities in Oriente Province and from Marianao in Habana 
Province collected entirely by H. Frere Clement in July 1935, and 
June, October, and December 1938. 

We were most fortunate in finding and rearing successfully the 
nymph of Microneura caligata. Full descriptions and figures of 
the nymph as well as adult male and female of this species, along 
with a description of the nymph of Neoneura maria (Scudder) are 
included in this paper. 

Collecting Localities in Las Villas Province 

1. Ponds and pools in a stream bed in Atkins Botanical Garden 
at Central Soledad near Cienfuegos. 

2. Stream six miles southeast of Soledad on Cienfuegos-Trin- 
idad road. 

3. Stream seven miles southeast of Soledad on Cienfuegos- 
Trinidad road. 

4. Brackish pond close to Cienfuegos-Trinidad road, near Trin- 
idad. 

5. Small, rocky stream in the Trinidad Mountains near Trini- 
dad, crossed by paved road to the Tuberculosis Sanitorium on 
top of the mountain. The stream is seven miles from the Cienfue- 
gos-Trinidad road, and is overgrown with dense vegetation, re- 
sembling a rain forest, especially above the road to a point where 



Westfall: Notes on Odonata of Cuba 69 

it issues from a cave in the side of the mountain. This small cave 
furnished a retreat during a heavy downpour. 

6. Large, rocky stream in the garden of the Tuberculosis Sani- 
tarium. The water here is excessively cold, so much so that the 
elaborate swimming pool built at the hospital could not be used. 

7. Small stream in Trinidad Mountains near Salto del Hana- 
banillo about 31 miles from Atkins Botanical Garden. 

8. Large shallow pond apparently formed by road construc- 
tion, about two miles beyond locality number seven. 

Suborder Anisoptera 

Family Gomphidae 

Aphylh caraiba (Selys). 2 $, 1 9, May 28-31, locality 1. 
Progomphus integer Hagen. 29$, May 29-June 7, localities 1, 3, 7; 
reared. 

Family Aeshnidae 

Aeshna psilus Calvert. 1 S , June 8, locality 5; reared. 

Anax amazili (Burmeister). [ANSP: 1 9 , June 1938, Marianao, Habana, 
H. Frere Clement]. 

Coryphaeschna adnexa (Hagen). 3 $ , May 29-June 7, locality 1; reared 
(emerged June 13 in Florida). 

Gynacantha ereagris Gundlach. [ANSP: 1 9 , July 1935, Loma del Gato, 
H. Frere Clement]. 

Family Libellulidae 

Brachymesia furcata (Hagen). 2 $ , 19, June 5, locality 4. 

Cannacria herbida (Gundlach). 8 $ , May 28-June 3, locality 1; reared. 

Cannaphila insuhris funerea (Carpenter). 20 S , 2 9 , May 29-June 7, 
localities 1 and 6. 

Dythemis rufinervis (Burmeister). 37 $ , 10 9 , May 29-June 7, localities 
1, 3, 6, and 7; [USNM: 19, August 19, 1946, El Caney, C. T. Ramsden; 
ANSP: 1 $ , 1 9 , July 1935, Loma del Gato and 1 9 , December 1938, Rente, 
Santiago Bay, H. Frere Clement]. 

Erythemis attala (Selys). 4 £, 1 9, June 3-6, locality 1. 

Erythemis haematogastra (Burmeister). 7 $ , May 28-June 6, localities 
1 and 3. This is a new record for the island. 

Erythemis plebeja (Burmeister). 30 $ , 12 9 , May 28-June 7, localities 
1 and 4; reared (nymphs were common). 

Erythemis simplicicollis (Say). 11 <2 , 4 9, May 28-June 6, localities 
1 and 4; [USNM: 1 $,3 9, May 27, 1904, Cayamas, E. A. Schwarz]. 

Erythrodiplax berenice subspecies? [ANSP: 3 9 , October and December 
1938, Rente, Santiago Bay, H. Frere Clement]. 

Erythrodiplax connata connata (Burmeister). [ANSP: 1 $ , 1 9 , July 
1935, Loma del Gato, H. Frere Clement]. Borror (1942, p. 178) lists a male 



70 Quarterly Journal of the Florida Academy of Sciences 

and a female from this same lot of specimens in the Cornell University collec- 
tion as connata connata. Further study of the connata group in the Antilles is 
needed to determine the relationship of justiniana with these almost clear- 
winged specimens and the form fraterna of Hagen. 

Erythrodiplax fervida (Erichson). 25 S, 4 9, May 28-June 6, locality 1. 

Erythrodiplax justiniana (Selys). 23 $ , 3 9 , May 29-June 7, localities 
1 and 4; [USNM: 5 S , 1 9, May 6-June 4, 1904 ; Cayamas, E. A. Schwarz]; 
reared. Wings of these individuals bore typical basal spot characteristic of this 
form. 

Erythrodiplax umbrata (Linnaeus). 3 <J , 2 9 , May 29-June 4, localities 
1 and 7; [USNM: 4 3,5 9 , May 27-June 2, 1904, Cayamas, E. A. Schwarz; 
1 9 , August 19, 1946, El Caney, E. T. Ramsden; ANSP: 2 3,1 9 , July 1935. 
Loma del Gato, H. Frere Clement; 13 $ , 18 9 , October and December 1938, 
Rente, Santiago Bay, H. Frere Clement]. 

Idiataphe cubensis (Scudder). 1 $ , June 5, locality 4; exuviae also col- 
lected. 

Lepthemis vesiculosa (Fabricius). 1 $ , 1 9 , June 5-6, localities 1 and 4; 
[ANSP: 2 $,S 9, December 1938, Rente, Santiago Bay, H. Frere Clement]. 

Libellula needhami Westfall. 1 9, May 31, locality 1; [USNM: 1$, 
5 9, May 27-June 2, 1904, Cayamas, E. A. Schwarz]. 

Macrodiplax balteata (Hagen). 2 $ , June 5, locality 4. 

Macrothemis celeno (Selys). 28 $ , 6 9 , May 29-June 7, localities 1, 
2, 3, and 6; [USNM: 2 $ , 1 9 , August 19-November 3, 1946, El Caney and 
Vista Alegre, Santiago, E. T. Ramsden; ANSP: 1 $ , 1 9 , July 1935, Loma del 
Gato, and 1 9, December 1938, Rente, Santiago Bay, H. Frere Clement]. 

Miathyria marcella (Selys). 1 9, June 1, locality 1; [ANSP: 1 9, Decem- 
ber 1938, Rente, Santiago Bay, H. Frere Clement]. The specimen we captured 
was flying very high along with other species, but we were able to shoot it 
with a pistol and dust shot. 

Micrathyria aequalis (Hagen). 38 £ , 9 9 , May 28-June 7, localities 1, 
2, 4, and 7; reared. 

Micrathyria debilis (Hagen). [USNM: 1$, May 30, 1904, Cayamas, 
E. A. Schwarz]. 

Micrathyria didyma (Selys). 9 $, 3 9 , May 29-June 7, locality 1; 
[USNM: 1 $, June 11, 1904, Cayamas, E. A. Schwarz]. 

Micrathyria dissocians Calvert. 15 $ , 3 9 , May 28-June 6, localities 1 
and 4; reared. 

Micrathyria hageni Kirby. 41 $ , May 29-June 7, locality 1; [ANSP: 
1 $, 1 9, December 1938, Rente, Santiago Bay, H. Frere Clement]. 

Orthemis ferruginea (Fabricius). 10 $ , May 29-June 7, localities 1 and 
3; reared; [ANSP: 2 $ , July 1935, Loma del Gato, and 2 $, December 1938, 
Rente, Santiago Bay, H. Frere Clement]. 

Pantala flavescens (Fabricius). 3 $ , I 9 , June 4, locality 8; reared; 
[USNM: 5 $,4 9, June 18-20, 1946, Vista Alegre, Santiago de Cuba, C. T. 
Ramsden; ANSP: 1 $ , July 1935, Loma del Gato, and 1 6 , December 1938, 
Rente, Santiago Bay, H. Frere Clement]. 



Westfall: Notes on Odonata of Cuba 71 

Pantala hymenaea (Say). [ANSP: 1 $ , June 1938, Marianao, Habana, 
H. Frere Clement]. 

Perithemis metella (Selys). 18 $ , May 28-June 7, localities 1 and 7; 
reared; [USNM: 1 $, May 27, 1904, Cayamas, E. A. Schwarz]. Racenis 
(1958) in a footnote states that Dr. K. Buchholz, who is doing a revision of 
the genus Perithemis, wrote to him that P. mooma Kirby 1889 is a synonym 
for P. domitia Drury 1773. Consequently, the species which has gone under 
'the name of domitia Drury in the revision of Ris (1930), in the manual of 
Needham and Westfall (1955), and elsewhere is to be known as P. metella 
(Selys) 1857. The species which has been referred to as mooma Kirby by Ris, 
Whitehouse (1943), and others should now be domitia Drury. In Arizona, at 
the San Bernardino Ranch in Cochise County, I collected numerous specimens 
of metella in July 1958. I found in separating metella from domitia and tenera 
that a character more constant than the presence or absence of crossveins in 
the subtriangle was the presence of three (metella) or two (domitia and tenera) 
paranal cells bordering the inner side of the subtriangle in the front wing. 

Scapanea frontalis (Burmeister). 24 $ , 2 9 , May 30-June 8, localities 3, 
5, and 6. [ANSP: 4 $ , 1 9 , July 1935, Loma del Gato, and 1 $ , December 
1938, Rente, Santiago Bay, and 2 $ , 1 9 , June 1938, Marianao near Habana, 
H. Frere Clement]. 

Tauriphila argo (Hagen). 1 $, June 1, locality 1. Dow (1931) listed a 
male from Soledad which he did not definitely assign to species. The speci- 
men here reported is probably the same species. It agrees quite well with 
specimens from Panama, Guatemala, and Colombia which I consider to be 
argo. 

Tauriphila australis (Hagen). 2 $ , 3 9 , May 29-June 6, locality 1. 

Tramea abdominalis (Rambur). 3 $ , May 29-June 1, locality 1. [USNM: 
1 9 , June 4, 1904, Cayamas, E. A. Schwarz; ANSP: 2 $ , December 1938, 
Rente, Santiago Bay, H. Frere Clement]. 

Tramea hinotata (Rambur). 3 $ , 4 9 , May 29-June 5, localities 1, 4, 
and 8. [USNM: 1 $ , May 27, 1904, Cayamas, E. A. Schwarz; ANSP: 2 $ , 
December 1938, Rente, Santiago Bay, H. Frere Clement]. 

Suborder Zygoptera 
Family Coenagrionidae 

Argiallagma minutum (Selys). 1 $ , 1 9, June 7, locality 1. These 
were taken in tandem and were the only ones seen. Geiiskes in 1943 de- 
scribed the nymph of this species and showed that it was not closely related 
to Argia as the generic name suggests, and as had been supposed by a number 
of authors. Indeed it seems to be much closer to Nehalennia. The long tibial 
spines used as the principal generic character for Argiallagma are not much 
longer than those of N. pallidula Calvert. Because of the long spines of palli- 
dula this species has often been determined as Argiallagma minutum in col- 
lections from Florida. All such specimens I have seen from Florida, including 
those of Byers, who added minutum to the Florida list in 1930, must be re- 
ferred to pallidula, and the species minutum should be deleted from the Flor- 



72 Quarterly Journal of the Florida Academy of Sciences 

ida list unless actual specimens can be found. It has been questionable wheth- 
er the genus Argiallagma should be retained, but supposed nymphs of pallidula 
recently taken in Florida are very similar to those of minutum and seem to 
be separable from nymphs of Nehalennia proper. At the present time I am 
proposing that we move pallidula to Argiallagma and retain the genus for these 
two known species. The adults are separable from the species of Nehalennia 
by the possession of a blackish metallic coloration instead of a brilliant green 
metallic color. The males of the two species are rather difficult to separate 
from each other, but the females show marked differences in the shape of the 
hind margin of the posterior lobe of the prothorax. 

Ceratura capreola (Hagen). 9 $, 49, May 29-June 7, locality 1; reared. 

Enallagma civile (Hagen). 5 $ , 3 9 , June 2-5, localities 2, 4, and 7. 
Specimens of this species in several collections have been found labeled 
Enallagma doubledaiji (Selys). Selys (1857, p. 469) listed doubledayi from 
Cuba based on a male from Latreille's collection. Gundlach (1888, p. 230) 
referred to this, but he did not collect the species in Cuba. Hagen (1861) 
made reference to Selys' paper but did not list the species from Cuba. Need- 
ham (1941, p. 14) said he had not collected doubledayi in Cuba and the only 
specimens he had seen from that source were "a few that were collected at 
'Placelas, Santa Clara' (Placetas, Las Villas) Province, Cuba by Dr. J. 
Acuna. ..." I have examined a male and female in alcohol from the Cornell 
University Collection labeled Enallagma doubledayi, June 28, 1940, Placelas, 
Cuba, by Dr. Needham. The appendages and penis of the male are almost 
identical with those of the Florida specimens, but the color pattern of the 
abdomen is strikingly different. In the Cuban male, segments 3 to 7 are black 
on the dorsum except for a very narrow basal ring. Segments 3 to 6 of the 
doubledayi from Florida are blue except for the apical sixth to fourth which 
is black. The female is somewhat teneral, but shows segment 8 to be entirely 
blue, whereas the female of doubledayi from Florida has the dorsum of this 
segment entirely black. The Cuban specimens are markedly smaller: male, 
length 25 mm., abdomen 21 mm., hind wing 13 mm.; female, length 27.8 
mm., abdomen 22 mm., front wing 17 mm. (hind wings incomplete). A 
search for more specimens of this species should be made to ascertain its true 
status. My present opinion is that it is distinct from doubledayi, but whether 
this is the same as the male in Latreille's collection reported by Selys is not 
certain as the description is so brief. The size reported by Selys ("Long., 
30-31; alae inf. 17-18; lat. alae inf. 3% millim.") seems to be more like the 
size of the Florida doubledayi than the Placetas male. However, since he 
reports only one male from Cuba in Latreille's collection these measurements 
could not have been taken from that specimen alone, but might in reality 
have been taken from Florida specimens. 

Enallagma cardenium Hagen. 38 $ ,25 9 , May 28-June 8, localities 
1, 2, 3, and 6; reared. The relationship of coecum and cardenium in Cuba 
is still confused and for the present I am considering all of my specimens as 
cardenium. 

Ischnura ramburi (Selys). 7 $ , 9 9 , May 28-June 8, localities 1, 3, 7, 
and 8; reared; [USNM: 1 9, May 8, 1904, Cayamas, E. A. Schwarz]. These 



Westfall: Notes on Odonata of Cuba 73 

specimens would probably fall into what has been called in the past the sub- 
species credula; however, they vary considerably as to the extent of black on 
the dorsum of segment 9 in the males, the character which has been used to 
separate these forms. 

Neoerythromma cultellatum (Hagen). 14 $ , 7 9 , May 28- June 2, lo- 
cality 1; reared. 

Telebasis dominicana (Selys). 41 $ , 14 9 , May 28- June 7, localities 1, 
2, and 7; reared; [ANSP: 1 9 , December 1938, Rente, Santiago Bay, H. Frere 
Clement]. 

Family Pseudolestidae 

This name used by Fraser (1957) to include Hypolestes is admittedly 
unsatisfactory as he includes in it a number of genera which do not form a 
natural group. It has been suggested by Dr. B. E. Montgomery in a letter to 
me that a separate family be used for Hypolestes and I believe this would be 
a partial solution to the problem. 

Hypolestes trinitatis Gundlach. 54 $ , 32 9 , May 30- June 8, locality 5; 
reared; [ANSP: 8 $,S 9, July 1935, Loma del Gato, E. A. Schwarz]. 

Family Lestidae 
Lestes tenuatus Rambur. 4 $ , 3 9 , June 3-4, localities 1 and 8; reared. 

Family Protoneuridae 

Microneura caligata Selys. 25 $ , 34 9 , May 30-June 8, locality 5; 
reared. Also ten nymphs were found in the Cornell University collection 
with a label in Dr. J. G. Needham's handwriting, "Neoneura sp. ? Rio San 
Antonio by Charco Azul, Trinidad Mts. Cuba 23 III '39". 

Neoneura carnatica Selys. [ANSP: 5 $ , June 1938, Marianao, Habana, 
H. Frere Clement]. 

Neoneura maria (Scudder). 34 $ , 15 9 , May 30-June 8, localities 3, 5, 6, 
and 7; reared. 

Protoneura capillaris (Rambur). 18 $ , 5 9 , May 29- June 7, locality 1. 
On June 7 a female, attended continuously by a male grasping her in the usual 
manner, was observed ovipositing in a small trickle of water between two 
pools of water in the stream bed at Soledad. This water had not connected 
the pools a few days earlier but rain had raised the water level. The female 
pressed the end of her abdomen against a dead palm leaf only about one-half 
inch below the surface of the water and then moved to repeat this, each time 
perhaps depositing a single egg. A search for nymphs here was unsuccessful. 

Life History Notes 

Microneura caligata Selys 

This is a beautiful little damselny which I found only at one 
stream in the Trinidad Mountains of Cuba. Dr. Needham had 
mentioned a single incomplete and teneral male at the Museum 



74 Quarterly Journal of the Florida Academy of Sciences 

of Comparative Zoology at Harvard, the only one he had seen. 
This specimen I studied in 1959. It had Hagen's pin labels on 
it and agreed with my specimens. There was also a second male 
with the data, "Mina Carlota, Trinidad Mts. 11-16 July, 1939, C. T. 
Parson". Microneura caligata was first mentioned in print by 
Hagen in 1867, without a description. Selys (1886) gave Hagen's 
name after the species, referring to the 1867 paper, but he did 
not state that the description had been submitted by Hagen so 
I am considering Selys as the first describer. 







Fig. 1. Microneura caligata. A, dorsal view of end of male abdomen; 
B, lateral view of same; C, ventral view of same; D, lateral view of end of 
female abdomen. 



Most of the M. caligata specimens I collected in Cuba were 
mating pairs, and were found along the very densely shaded stream 
mentioned earlier. In a small pool, just outside the cave from 
which this stream issued, I found two exuviae on a rock about 
two inches above the water line. Other rocks were picked up, 
and large nymphs were discovered clinging to the under sides. 
These were reared at the laboratory. Nymphs of Hypolestes trini- 
tatis were also taken from the under sides of rocks in deeper pools 



\ 



\ 



/ 



/ 



> 



N>\ 



>- 
A 



| ; _ / >■> 



., 



• : 



/- 



< 



Fig. 2. Microneura caligata. Dorsal view of nymph, labium, and external 
view of left lateral gill. 



76 Quarterly Journal of the Florida Academy of Sciences 

of the same stream and reared. One pair of M. caligata was ob- 
served ovipositing. The male continued to clasp the female in 
the usual manner as she oviposited. She lit on a plant close to a 
moss-covered rock, which was partly submerged, and oviposition 
occurred on this rock just below the surface of the water. In dense 
shade, where this species occurred, the thick white pruinosity of 
the legs of the male was quite conspicuous. 

Adult Male: The head and thorax are rather robust compared 
with other members of the Protoneuridae. Labrum, postclypeus, 
dorsal surface of the head posterior to the antennae, and the rear 
of eyes black with metallic blue and violet reflections. Antecly- 
peus obscure yellowish. Genae, frons anterior to the antennae, 
and extreme base of antennae reddish orange. The light stripe 
anterior to the antennae is sometimes divided in the middle by 
the black. Prothorax with median and posterior lobes chiefly red- 
dish orange, the anterior and lateral lobes mostly black. Dorsum 
of thorax black with metallic reflections, this black extending ven- 
trally on to the metepimeron. Narrow humeral stripe of reddish 
orange complete and extended to cover the outer corner of meso- 
stigmal plate. A yellowish stripe, wider than the humeral, covers 
the spiracle and extends upward about two-thirds the length of 
the metepisternum. Ventro-lateral border of metepimeron broadly 
yellowish. Dark markings on the venter of thorax. 

Legs long, with coxae and proximal half of femora reddish 
orange; coxae of 2nd and 3rd legs with a large black spot laterally. 
Femora noticeably compressed and expanded at distal end, the 
distal half black and covered with a dense white pruinosity which 
may add to the apparent width of the femora. Tibiae yellowish, 
externally; blackish internally; tarsi mostly dark. Femora and 
tibiae each with 2 rows of long spines evenly spaced, the metathor- 
acic femora with 10-12 in the external row, the tibiae with 8-9. 

Wings hyaline, nodus not strongly oblique, the subcosta ending 
distal to the origin of M 3 . 

Abdominal segments 1 and 2 black dorsally, some yellow on 
latero-ventral margin. Segments 3 to 7 black with bright blue 
basal spots on 3 about 5/6 the length of segment, on 4 about 1/3, 
on 5 about 1/5, on 6 about 1/6, on 7 about 1/15, divided in the 
middle; segments 8 and 9 almost entirely blue on the dorsum, 
latero-ventral margins narrowly black; segment 10 all black, its 
hind margin with a shallow notch dorsally. 



Westfall: Notes on Odonata of Cuba 77 

Appendages brown to black, in length about equal to segment 
10, the simple inferiors slightly longer than the complicated su- 
periors, which have heavily chitinized outer portion and a heavily 
chitinized medial branch which extends ventrally. There is a 
thinner, less chitinized curved fringe as illustrated in figure 1, 
A-C. 

Adult Female: The reddish orange markings of the male are 
replaced by yellow. Light markings of prothorax of male reduced, 
so that almost entire prothorax is black, small spot on side of 
median lobe and one on lateral extension of posterior lobe. Meso- 
stigmal plates extremely complicated and difficult of description, 
projecting strongly dorsad. Thoracic light markings as in the 
male but the humeral stripe is interrupted near the posterior end 
and sometimes widely in the middle also, thus forming three iso- 
lated short streaks. 

Abdomen dorsally is black with metallic reflections, only the 
slightest indication of yellow basal rings on 3 to 7 which are inter- 
rupted middorsally. Segment 8 has the latero-ventral margin 
widely yellow and 9 has a large yellow spot laterally which reaches 
the ventral margin. Ovipositor strongly marked with yellow dor- 
sally, brown ventrally, the black styles extending beyond end of 
abdomen. Appendages short, about equal to length of segment 
10 at midlateral height. End of abdomen is shown in figure 1,D. 

Nymph: A light tan nymph with inconspicuous color pattern. 
Head nearly five-eighths as long as wide, obscurely patterned. Eyes 
noticeably protruding from sides of head and with about 13 alter- 
nating light and dark dorso-ventral bands. Caudo-lateral angles 
of head rounded, provided with several stout setae. Hind margin 
excavated. Antennae longer than the head which is about 2.25 
mm.; dark brown, lighter at joints, with apical one-half of sixth 
segment and all of seventh white. Bases of antennae separated 
by slightly more than 1 mm. Length of seven segments in ratio 
of 9:14:28:25:14:9:5. 

Lateral margins of pronotum dark brown. Legs long, hind 
femora reaching slightly beyond tips of wing pads and almost to 
apex of abdominal segment 6 when extended alongside. Legs 
slightly flattened as in adult, feebly banded; tibiae with only a few 
almost invisible hairs. Wing pads reaching to middle of segment 
5 or 6. 



78 Quarterly Journal of the Florida Academy of Sciences 

Abdomen tan with three longitudinal dorsal light bands which 
may be interrupted, also a light lateral band extending to the lat- 
eral carinae. Surface of abdomen almost devoid of setae except 
on the posterior margins and ventral surfaces of the end segments. 
Ovipositor of female nymphs extending well beyond apex of seg- 
ment 10. 

Caudal gills about equal to length of abdomen; median gill 
slightly shorter than laterals. Basal three-fifths of each gill thicker 
and darker than the apical portion; this darker portion of lateral 
gill slightly greater in extent on the ventral margin than on the 
dorsal. Lateral gills with a strong lateral keel provided with 
about 30 short stout setae along the length of the thickened basal 
portion; dorsal setae about 20, ventral setae about 30, the last 5-6 
just before the thinner portion of gills much farther apart than 
others; apical part white, slightly grayish near base, with long 
hairs on margins. Median gill with about 30 setae on ventral mar- 
gin, 25 on dorsal margin, the last 4-5 on dorsal margin larger and 
farther apart. 

Labial mask in folded position reaching level of mesocoxae. 
Prementum of labium about five-sixths as wide as long, basal width 
one-third of maximum width; the median lobe strongly convex 
and minutely denticulate; setae of lateral margin large, about 
15-19 on slightly more than the distal half; irregular group of 4-9 
small setae on the dorsal surface, each side of the midline, about 
a third the distance from base to apex, besides the one large pre- 
mental seta each side of the midline anteriorly. Palpal lobes long, 
movable hook as long as portion of lobe basal to it. Distal margin 
of each palpal lobe with a long incurved mesal hook (end hook) 
and a shorter pointed lateral hook which is about one-third as 
long as the movable hook; palpal setae (lateral setae) three each 
side even in half grown nymphs (two of 21 nymphs have four 
palpal setae on one side). 

Total length 16 mm., abdomen 6 mm., lateral caudal gills 5.6 
mm., hind femora 3.12 mm., antennae 2.5 mm. Described from 
eight nymphs in last instar, six exuviae (two reared in laboratory, 
two taken in transformation, and two collected with nymphs in 
field), and seven half-grown nymphs. Measurements taken from 
average single nymph in last instar. 



A 



% 



\ 

4. 



I 



W^x 



/S 



,-k 



A 






J4*- 



r 



Fig. 3. Neoneura maria. Dorsal view of nymph, labium, and external 
view of left lateral gill. 



80 Quarterly Journal of the Florida Academy of Sciences 

Neoneura maria (Scudder) 

This species was found commonly on several streams, pairs 
ovipositing on floating vegetation similar to Neoneura aaroni Cal- 
vert as observed in Texas. The nymphs were all collected from 
the under sides of rocks lifted from the stream. All of the adults 
were clear blue with no indication of the reddish coloration of 
all the specimens of carnatica I have seen. This color was char- 
acteristic of the one male reared and other young adults, as well 
as older specimens which were pairing and ovipositing. No car- 
natica were found in the areas where maria occurred. 

Nymph: A tan nymph with rather definite color pattern of 
dark brown, legs conspicuously banded. Head noticeably pat- 
terned; two squarish light-colored areas on frontal shelf anterior 
to antennal bases; conspicuous rounded brown spot each side of 
midline near excavation at hind margin. Head about five-sevenths 
as long as wide. Eyes not noticeably protruding from sides of 
head (as they do in Microneura caligata) and uniformly colored. 
Caudo-lateral angles of head rounded, with several stout setae. 
Antennae equal to or slightly shorter than length of head; two 
basal segments light tan, 3-6 dark brown with increasing amount 
of white at apices of segments, 7 all white. Length of segments in 
ratio of 9:11:21:15:9:6:4. Bases of antennae separated by less than 
1 mm. 

Lateral margins of pronotum dark brown. Legs shorter than 
in Microneura caligata, hind femora not reaching to end of wing 
pads, about to middle of abdominal segment 5 or less. Wing pads 
just reaching segment 6, or in more extended specimens to about 
one-fourth length of segment 5. Femora with three conspicuous 
dark bands, tibiae with only one. Femora with scattered stout 
setae; tibiae with thick brush of long hairs on outer side. 

Abdomen with two conspicuous dark bands on dorsum, some- 
times merging in the midline. Caudo-lateral angles of basal seg- 
ments at lateral carinae light, remainder of lateral margins dark. 
Dorsal surface of abdomen with many more stout setae than in 
Microneura caligata. Ovipositor of female nymph extending be- 
yond apex of segment 10. Caudal gills much shorter than abdo- 
men, median gill slightly shorter than lateral. Basal portion of 
gills thicker and darker than apical part; the distal margin of the 
darker area in lateral gills diagonal, reaching to three-fifths length 



Westfall: Notes on Odonata of Cuba 81 

of gill on ventral margin, to less than half on the dorsal margin. 
The lighter distal area with dark markings more pronounced in 
some specimens than others, sometimes appearing as three or four 
quite separate bands, or a single diffuse band as shown in figure 3; 
apex of gills more attenuate with longer tips in some specimens 
than as shown in drawing. Lateral gills with a strong lateral keel 
provided with about 25-30 short stout setae along the length of 
the thickened basal portion; ventral setae about 28-35, the more 
distal setae much enlarged; dorsal setae about 10-12, the distal 
ones not much larger than the basal ones, all of them more irreg- 
ular in position than the ventral setae. Median gill with the basal 
thickened area about equal on the two margins; setae about equal 
in number, 12-15, the apical ones much larger on both margins. 

Labial mask in folded position not reaching level of mesocoxae. 
Prementum of labium more than nine-tenths as wide as long, the 
median lobe strongly convex and minutely denticulate; basal width 
slightly more than one-third the maximum width; setae of lateral 
margin large, about 12-16, on about the distal half; setae on the 
dorsal surface near base much less evident and fewer in number 
than in Microneura caligata, sometimes appearing quite absent; 
one large premental seta each side of the midline, even in small 
nymphs with wing pads barely visible, though the smallest nymphs, 
4.6 mm. long, seem to lack it. Palpal lobes long, movable hook 
about two-thirds as long as portion of lobe basal to it. Distal 
margin of each palpal lobe with a long incurved mesal hook (end 
hook) and a shorter pointed lateral hook which is about one-third 
as long as the movable hook; palpal setae (lateral setae) three 
each side even in half grown nymphs, four on one side in two 
specimens. Very small nymphs, 4.6 mm. long, have only one 
palpal seta; those 6.6 and 9.2 mm. long have two. 

Total length of full-grown nymphs 12.7-13.9 mm., abdomen 
5.5-6.1 mm., lateral caudal gills 3.5-3.7 mm., hind femora 2 mm., 
antennae 1:7 mm. Described from one reared specimen, one ex- 
uviae taken in field, four nymphs in last instar, and 36 smaller 
nymphs. 

Comparison: These nymphs are remarkably like those of car- 
natica which Needham described in 1939. I have his nymphs be- 
fore me and find it difficult to see distinguishing characters that 
are sharply defined. The adults of carnatica are larger and more 
robust in general than my specimens of maria and the same holds 



82 Quarterly Journal of the Florida Academy of Sciences 

true of the nymphs. The setal counts of the labium are the same. 
Of 26 nymphs of carnatica, two have four palpal setae on both 
sides, all others have three. The gills seem to offer the best char- 
acters for separation, though I do not have very many gills of ma- 
ture nymphs of maria for statistical comparisons. The lateral con- 
strictions seem to be more noticeable between the thickened 
basal and thinner apical portions of gills in carnatica. The setal 
counts of gills in mature nymphs are considerably higher in car- 
natica, the lateral gills having about 40-45 on the ventral margin, 
maria 28-35. Similar proportions hold for the other setose surfaces 
of the gills, though with a larger number of mature nymphs of 
maria this difference might not be so pronounced. Dr. Needham 
described the nymphs of carnatica as blackish, but they must have 
faded considerably in alcohol for now they are about the color of 
maria nymphs which I took. These two species are certainly close 
in morphology of adult and nymph, despite the great color differ- 
ences in the adult males. I believe there are subtle differences in 
the appendages. Like Williamson (1917, pp. 236-239) I am still 
compelled to consider these two insects as distinct species after 
observing maria in the field and seeing no intermediates between 
it and carnatica. Further study of the series now available before 
us may reveal other constant morphological differences. 

Hypothetical List 

Several species of Zygoptera which have been reported by 
various authors in the past should, I believe, be dropped from the 
Cuban list until actual specimens are taken. 

Family Calopterygidae 

Hetaerina cruentata (Rambur). This was listed by Selys and 
Gundlach, but no specimens from Cuba were cited, nor have any 
been found to date. 

Family Coenagrionidae 

Amphiagrion saucium (Burmeister). The record of this species 
seems to be based on a female from Latreille's collection, listed 
uncertainly by Selys as Agrion discolor? Burmeister. He stated 
that it might possibly be the female of Agrion dominicanum Selys. 



Westfall: Notes on Odonata of Cuba 83 

In 1876 Selys placed discolor as a synonym of Ampkiagrion saucium 
(as suggested by Hagen in 1861). In addition to the fact that Selys 
was uncertain of the identity of Latreille's specimen, no further 
specimens from Cuba have been reported that could be assigned 
to A. saucium. It is very likely that Latreille's specimen was not 
saucium and this species should therefore be omitted from the 
Cuban list. 

Diceratobasis macrogastcr (Selys). This has been included in 
lists of the Odonata of Cuba under several different genera, in- 
cluding Agrion, Nehalennia, and Telebasis. Selys (1857) clearly 
listed it as only possible in Cuba, citing it as from Jamaica. Gund- 
lach included it in his list, also, but no actual specimens were re- 
ported from Cuba. I first saw nymphs which I supposed to be this 
species taken from bromeliads in Jamaica in 1952 by Dr. A. M. 
Laessle. After I had collected and reared them myself in Jamaica 
in the summer of 1960, I gave Dr. Laessle the definite determina- 
tion for his paper (1961). The nymph is so unique that I have no 
reservation in accepting for it Kennedy's (1920) genus, Dicerato- 
basis, and the nymphal description will appear soon in a paper 
now in preparation. There is no reason to include it in the list 
of Cuban species until specimens are taken there. So far as is 
known, it is endemic to Jamaica. 

Telebasis vulnerata (Hagen). It is questionable whether this 
species has ever been taken in Cuba, despite the fact that Hagen 
(1861, p. 86) listed it as from "Cuba (Poey)" in his original descrip- 
tion. It is possible that Gundlach later misidentified his specimens 
because he interpreted Hagen's description of the male append- 
ages of vulnerata as being from the lateral view, whereas they were 
described from the dorsal view. Gundlach said vulnerata lived all 
over the island of Cuba, but that he had not observed dominicana. 
Collectors today find dominicana commonly, but never vulnerata. 
I have again studied types of these species at the Museum of Com- 
parative Zoology at Harvard and we are associating them cor- 
rectly. 

Acknowledgments 

The work here reported was made possible by a grant from 
the National Science Foundation. My wife, who accompanied 
me to Cuba, assisted also in some of the details of preparing this 
paper for publication. Dr. and Mrs. Duncan Clement and mem- 



84 Quarterly Journal of the Florida Academy of Sciences 

bers of their staff at the Botanical Garden in Soledad cooperated 
to make our stay there most successful. The drawings are the 
work of Esther Coogle, former staff artist for the Department of 
Biology, University of Florida. 

Literature Cited 

Borror, Donald J. 1942. A revision of the Libelluline genus Erythrodiplax. 
Ohio State Univ., Biol. Series, no. 4, pp. 1-286, pis. 1-41. 

Byers, C. Francis. 1930. A contribution to the knowledge of Florida Odo- 
nata. Univ. of Florida Press, Gainesville, 327 pp. 

Calvert, Philip P. 1919. Gundlach's work on the Odonata of Cuba: a 
critical study. Trans. Amer. Ent. Soc, vol. 14, pp. 335-396. 

Dow, Richard. 1931. Odonata from Santa Clara, Cuba. Proc. Biol. Soc. 
Wash., vol. 44, pp. 55-60. figs. 

Fraser, F. C. 1957. A reclassification of the order Odonata. Royal Zool. 
Soc. New South Wales, 133 pp. 

Geijskes, D. C. 1943. Notes on Odonata of Surinam. IV. Nine new or 
little known zygopterous nymphs from the inland waters. Annals Ent. 
Soc. Amer., vol. 36, pp. 165-184. 

Gundlach, Juan. 1888-1890. Contribution a la entomologia cubana. Vol. 
2, pt. 3, Neuropteros Havana, pp. 189-281. 

Hagen, Hermann. 1861. Synopsis of the Neuroptera of North America. 
Smithsonian Misc. Collections, Washington, xx + 347 pp. 

. 1867. The Odonat-fauna of the island of Cuba. Proc. Boston Soc. 

Nat. Hist., vol. 11, pp. 289-294. 

Kennedy, Clarence H. 1920. Forty-two hitherto unrecognized genera and 
subgenera of Zygoptera. Ohio Jour. Sci., vol. 21, pp. 83-88. 

Laessle, Albert M. 1961. A micro-limnological study of Jamaican bro- 
meliads. Ecology, vol. 42, no. 3, pp. 499-517. 

Needham, James G. 1939. Nymph of the Protoneurine genus Neoneura 
(Odonata). Ent. News, vol. 50, pp. 241-245, figs. 1-6. 

. 1941. Life history notes on some West Indian Coenagrionine 

dragonflies (Odonata). Jour. Agric. Univ. Puerto Rico, vol. 25, pp. 1- 
18, figs. 1-5. 

Needham, James G., and Minter J. Westfall, Jr. 1955. A manual of the 
dragonflies of North America. Univ. of Calif. Press, Berkeley, xii + 
615 pp. 



Westfall: Notes on Odonata of Cuba 85 

Racenis, J. 1958. Los odonatos neotropicales en la coleccion de la Facultad 
de Agronomia de la Universidad Central de Venezuela. Acta Biol. 
Ven., vol. 2, art. 19, pp. 179-226. 

Ris, F. 1930. A revision of the Libelluline genus Perithemis (Odonata). 
Misc. Pub. Mus. Zool. Univ. Mich., vol. 21, pp. 1-50, pis. 1-9. 

Selys-Longchamps, Edmond de. 1857. Odonates de Cuba. In R. de la 
Sagra, Histoire physique, politique et naturelle de File de Cuba. 
Paris, pp. 436-472. 

. 1886. Revision du Synopsis des Agrionines. Premiere partie. Mem. 

Acad. Roy. Belg., vol. 38, pp. i-iv, 1-233. 

Whitehouse, Francis C. 1943. A guide to the study of dragonflies of 
Jamaica. Bull. Inst, of Jamaica, Science Ser., no. 3, pp. 1-67. figs. 

Williamson, E. B. 1917. The genus Neoneura (Odonata). Trans. Amer. 
Ent. Soc, vol. 43, pp. 211-246. 

Department of Biology and Florida State Museum, University 
of Florida, Gainesville, Florida. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



DIETARY VITAMIN A AND COPPER CONCENTRATION 
IN THE LIVER AND HEART OF RATS 

Marie L. Kraemer, M. C. Jayaswal, J. F. Easley, 
and R. L. Shirley 

Copper and vitamin A are interrelated in a number of ways. 
Copper may destroy vitamin A in feedstuffs (Bhattacharya and 
Basu, 1954; Halverson and Hendrick, 1955). Balakhovskii and 
Drozdova (1957) related keratinization of epithelial tissue to copper 
catalysis in the absence of the inhibiting function of carotenoids. 
Kramer and Tarjan (1960) fed approximately 100, 500 and 1500 
ppm of copper and 15 International Units (I.U.) of vitamin A per 
day to rats in a basal diet containing 13 ppm of copper and reported 
that the vitamin A was lower in liver of those supplemented with 
copper. Fifteen mg. of copper per day was required to increase 
the copper in the liver. Shirley et al. (1962) observed that vitamin 
A supplementation decreased the deposition of copper in the liver 
and copper supplementation slightly but significantly increased 
(P < .05) the level of vitamin A in the liver of swine. 

The experiment reported here was designed to study further 
this interrelation of dietary copper and vitamin A on their concen- 
tration in the liver and heart of rats. 

Experimental 

Forty-eight male rats, 26 days of age, were given a basal diet 
that contained the following ingredients, in percentages: skim milk 
powder (Borden's non-fat), 49.0; sucrose, 48.5; pork lard, 2.0; so- 
dium chloride, 0.5; ferrous sulfate, 0.002; manganous sulfate, 0.001; 
and thiamine. HC1, 0.0003. By analysis this diet contained 6 ppm 
copper and was fed to half the rats as prepared. Copper sulfate 
was added to part of the diet in order that half of the rats were 
fed 56 ppm of copper. Twelve rats of each dietary copper group 
were given orally by a medicine dropper 200 I.U. of vitamin A 
acetate every other day. This provided 4 treatments of a factorial 
design in which each level of dietary copper was fed with and 
without added vitamin A. 

The rats were fed for 90 days, after which time they were sacri- 
ficed. Liver and heart were removed, frozen, and analyzed for 



Kraemer et al.: Vitamin A and Copper in Rat Tissues 



87 



O < 

3 > 

so c 



VITAMIN A 



I.U./GM, 



FRESH 

ro 

O 

i 


WEIGHT 
o o 

1 1 




COPPER 



P.RM. DRY WEIGHT 




Fig. 1. Effect of dietary copper and vitamin A levels on the concentra- 
tion of these nutrients in the liver of rats. 



88 Quarterly Journal of the Florida Academy of Sciences 

copper and vitamin A within a few days. Copper was determined 
using versenate according to Cheng and Bray (1953) and caro- 
tenoids and vitamin A by the method of Gallup and Hoefer (1946). 
Statistical analysis of the data was made as recommended by Snede- 
cor (1956). 

Results and Discussion 

Figure 1 graphs the data obtained for the effect of the treat- 
ments on the concentrations of vitamin A and copper in the liver. 
The rats supplemented with vitamin A contained approximately 
30 times as much vitamin A in the liver as those not supplemented. 
Both the supplemented and unsupplemented vitamin A dietary 
groups that were fed 56 ppm of copper had slightly more vitamin 
A present than those that were fed the 6 ppm of copper. This in- 
crease in vitamin A owing to dietary copper was not significant, 
in contrast with the results observed with swine (Shirley et al., 
1962). These observations are also in contrast with those of Kramer 
and Tarjan (1960), who found less vitamin A present in the liver 
of rats fed copper in the range of 100, 500, and 1500 ppm. This 
suggests that since vitamin A prevents copper from producing 
keratinization of tissues, copper in slight excess of dietary needs, 
as given in the present study, may stimulate deposition of the vita- 
min; but when greater excesses are given as reported by Kramer 
and Tarjan (1960), the copper then may decrease the vitamin A 
present by catalyzing its oxidation. 

As shown in figure 1, approximately 25 to 50 per cent more 
(P < .01) copper was found in the liver of rats fed 56 ppm than in 
those fed 6 ppm. These results are also different from those of 
Kramer and Tarjan (1960) which showed that 1500 ppm of dietary 
copper was required before there was an appreciable increase in 
copper concentration in the liver. This difference may be explained 
on the basis that Kramer and Tarjan had 18 per cent protein in 
their diets compared to approximately 13 per cent in the present 
study. McCall and Davis (1961) reported that increasing the casein 
protein dietary level from 10 to 17.5 per cent decreased the deposi- 
tion of radioactive and total dietary copper in the liver of rats. 

Table 1 presents the data obtained for the effects of the dietary 
treatments on the deposition of copper, carotenoids and vitamin 
A in the heart. In contrast to the liver, the 56 and 6 ppm of dietary 
copper resulted in only slight variations (8.3 to 11.1) in the amounts 



Kraemer et al.: Vitamin A and Copper in Rat Tissues 



89 



of the element deposited in the heart. Carotenoids calculated 
as ^-carotene ranged from 0.32 to 0.72 fxg. per gram wet weight in 
the hearts of the various dietary groups. Vitamin A was not de- 
tected in any of the hearts. 



TABLE 1 

Effect of dietary vitamin A and copper intake on 
these nutrients in the heart 



the concentration of 



Cu, ppm 


56 


6 


56 


6 


Vitamin A, I.U. per day 


100 


100 








Number rats 


11 


12 


11 


7 


Cu, ppm* 


8,3 


9.4 


10.2 


11.1 


^-carotene, /ng. 










per gm. wet wt. * 


0.68 


0.72 


0.32 


0.71 


Vitamin A 















Differences are not statistically significant. 



Acknowledgment 



This study was financed in part by a grant-in-aid from the 
National Heart Institute, P.H.S. (H-1318). 



Summary 

Male rats were fed, from 26 until 116 days of age, semi-purified 
diets with skim milk powder as the source of approximately 13 
per cent protein and contained 6 ppm of copper by analysis, or 56 
ppm copper by addition of copper sulfate. Rats fed these two 
diets were subdivided in such a manner that half of each copper 
dietary group were given 200 I.U. of vitamin A acetate in solution 
every other day. When sacrificed, liver and heart ventricle were 
analyzed for copper and vitamin A. With supplemental dietary 
copper and vitamin A these nutrients increased in the liver but not 
in the heart. Traces of carotenoids, but no vitamin A, were found 
in the heart. Vitamin A had no significant effect on the amount of 
copper deposited in the heart and liver. Copper supplementation 
had no effect on the amount of vitamin A in the heart and liver. 



90 Quarterly Journal of the Florida Academy of Sciences 

Literature Cited 

Balakhovskii, S. D., and N. N. Drozdova. 1957. The mechanism of action 
of vitamin A (retinol). The copper-polyene antagonism and keratiniza- 
tion of the epithelial tissue. Biokhimiya, vol. 22, pp. 330-335. Chem. 
Abstrs., vol. 51, pp. 11420. 

Bhattacharya, S., and U. P. Basu. 1954. Vitamin A in solution. VII. 
Effect of trace metals. Jour. Indian Chem. Soc, vol. 31, pp. 241-249. 

Cheng, K. L., and R. H. Bray. 1953. Two specific methods of determining 
copper in soil and plant material. Anal. Chem., vol. 25, pp. 655-659. 

Gallup, W. D., and J. A. Hoefer. 1946. Determination of vitamin A in 
liver. Ind. Eng. Chem., Anal. Ed., vol. 18, pp. 288-290. 

Halverson, A. W., and C. M. Hendrick. 1955. Effect of trace minerals and 
other dietary ingredients upon vitamin A stability in stored poultry diets. 
Poultry Science, vol. 34, pp. 355-360. 

Kramer, Magdalene, and R. Tarjan. 1960. Studien liber den Carotins- 
toffwechsel V. Die Wirkung von Kupfer auf die Verwertung von Caro- 
tin. Internat. Zeitschr. Vitaminforsch., vol. 30, pp. 271-278. 

McCall, J. T., and G. K. Davis. 1961. Effect of dietary protein and zinc on 
the absorption and liver deposition of radioactive and total copper. 
Jour. Nutrition, vol. 74, pp. 45-50. 

Shirley, R. L., T. N. Meacham, A. C. Warnick, H. D. Wallace, J. F. 
Easley, G. K. Davis, and T. J. Cunha. 1962. Gamma irradiation 
and interrelation of dietary vitamin A and copper on their deposition 
in the liver of swine. Jour. Nutrition, vol. 78, pp. 454-456. 

Snedecor, C. W. 1956. Statistical methods. The Iowa State University 
Press, Ames, Iowa, ed. 5. 

Department of Animal Science, University of Florida, Gaines- 
ville, Florida (present address of Marie L. Kraemer, Mercedes, 
Corrientes, Argentina; of M. C. Jayaswal, Marine Laboratory, St. 
Petersburg, Florida). 

Florida Agricultural Experiment Stations Journal Series No. 
1849. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



TEMPERATURES OF THREE BREEDS OF YEARLING 
STEERS IN SOUTH FLORIDA 

C. E. Haines and M. Koger 

The normal body temperature of beef cattle is considered to 
range between 98.0° and 102.4° F. Although Morrison (1957), 
Findlay (1950) and others agree that the normal rectal temperature 
of beef cattle should be about 101.0-101.5° F, a study by Haines 
(1961) showed that 104.0° F. was the average temperature for a 
group of Hereford yearlings in South Florida. Reports by Brody 
(1956), Bonsma (1948), Garrett et al. (1960), Findlay (1950), Rhoad 
(1938), and Rieck and Lee (1948) all point out that body tempera- 
tures of cattle are affected to some extent by air temperature and 
other climatic factors. These same reports also indicate that all 
breeds of cattle do not respond in the same manner to changes in 
atmospheric conditions. 

Most of the studies reported on heat tolerance of cattle have 
pertained to mature dairy cattle and only a few to beef cattle 
breeds. More recent studies have involved the use of controlled 
temperature and humidity chambers for determining the effects 
of these items on physiology and production items. Although 
these artificial conditions are useful for stabilizing environmental 
conditions for such studies, information obtained under normal 
environmental conditions is also of importance. 

It has been suspected that high humidity, intense radiation, and 
environmental temperatures in South Florida effect the body tem- 
peratures of cattle in this area. Therefore, the investigation re- 
ported herein involved recording the rectal temperatures of steers 
grazing on pasture in South Florida to verify this theory. 

Experimental 

Rectal temperatures of 12 yearling steers were recorded at 
two-week intervals for a complete year. The following year, 12 
different yearling steers were used for the same purpose. The 
steers were approximately one year old at the beginning of each 
year of the study. During each year, four Angus, four Brahman, 
and four Hereford steers contributed the data. All animals were 
born and raised in the experiment station herd, and their dams 



92 Quarterly Journal of the Florida Academy of Sciences 

had been members of the same herd for several years. Therefore, 
these animals were considered acclimated to the area. The steer 
calves and their dams had been maintained continuously on Rose- 
lawn St. Augustinegrass pastures. 

To secure rectal temperatures, the animals were confined in 
a covered chute approximately 15 to 30 minutes after walking in 
leisurely from nearby pastures. A Fahrenheit thermometer was 
inserted four inches into the rectum and held in the passage for 
at least three minutes. The actual air temperature was recorded 
at the beginning and end of each data collection period, by using 
the average of two thermometers located on opposite sides of a 
post. The midway point between the initial and final air tempera- 
ture was used as the air temperature for that particular collection 
period. The collection periods usually lasted from 8:30 AM to 
11:00 AM. 

Individual data were compiled by breed, year of study, and 
period of the year and subjected to an analysis of variance as de- 
scribed by Snedecor (1956). Correlation and regression coefficients 
for body temperatures and environmental temperatures were cal- 
culated also. 

Results and Discussion 

The overall average body temperature of all 24 steers used 
during the two year period was 103.9° F. The average rectal 
temperatures by breed were 104.2°, 103.4 c , and 104.1° F for 
Angus, Brahman, and Herefords, respectively, with the Brahmans 
differing significantly (P < .05) from the Angus and the Hereford. 
Although these temperatures seem high, reports by Rieck and Lee 
(1948), by Bonsma (1948), and the comments by Findlay (1950) 
and Morrison (1957) indicate that calves have higher temperatures 
than mature cattle. These researchers have shown that heat tol- 
erance is the lowest in calves and that it increases gradually up 
to about four years of age. Since the steers involved in the present 
study were between one and two years of age when contributing 
these data, their body temperatures might be expected to decline 
slightly as they reach maturity. However, data on such response 
was not obtained in this study. 

As air temperatures increased, the rectal temperatures rose 
also. This same trend was reported by Brody (1956). Findlay 
(1950) stated that a rise of 5 C in the body temperature of Angus 



Haines and Koger: Body Temperatures of Cattle 93 

cows could be expected at an air temperature of 100° F, and 
Bianca (1963) found that rectal temperatures of calves rose almost 
linearly with the length of exposure to high temperatures in en- 
vironmentally controlled chambers. The temperatures recorded 
in Table 1 show the rise occurring in body temperature of steers 
in this study as air temperatures increased. Air temperatures varied 
from 51 to 91° F at time of observations, and the values shown 
in this table were selected as representatives of the trend observed. 

Statistical analysis indicated that air temperatures had a sig- 
nificant effect (P < .01) on body temperature in each of the breeds 
in this study. 

Regression coefficients indicated that a rise of 10° F in air tem- 
perature caused a rise of 0.48° in the body temperature of Angus 
steers and an increase of 0.43° in the body temperature of Brahman 
and Hereford steers. The correlation coefficients between atmos- 
pheric temperature and body temperature were 0.61, 0.57, and 
0.70 for the Angus, Brahman and Hereford breeds, respectively. 
The overall correlation coefficient for breeds combined was 0.62. 

TABLE 1 

Environmental temperatures and average body temperatures of yearling steers 



Dates of 
Observation 






Air 
Temper- 
ature * 


Steers - 

per 
Breed 


Angus 


Breed of Steer 
Brahman 


Here- 
ford 


Feb. 9 






51 


4 


103.1 


101.6 


103.0 


Jan. 12, Mar. 9 






68 


8 


103.7 


102.7 


103.6 


Nov. 30, Dec. 29 


i 




71 


8 


103.4 


103.0 


103.4 


April 5, Nov. 2 






75 


8 


103.6 


103.0 


103.4 


Jan. 25, Feb. 8, 


April 


6 












April 19, Oct. 19 






79 


20 


104.2 


103.6 


104.0 


Mar. 22, June 1, 


Nov. 


16 


82 


12 


104.1 


104.0 


104.0 


May 4, Aug. 24, Sept. 1 




87 


12 


104.2 


103.5 


104.1 


May 31, June 28, 


Sept. 


6 


89 


12 


105.2 


104.1 


104.8 


July 13, Aug. 23 






91 


8 


105.0 


104.0 


104.8 



*A11 temperatures expressed in degrees Fahrenheit. 



94 Quarterly Journal of the Florida Academy of Sciences 

Summary 

Rectal temperatures of four Angus, Brahman and Hereford 
yearling steers were taken at two week intervals throughout the 
year. A similar group provided information for a second year of 
observations. All steers originated in the area and were main- 
tained on grass pasture. 

The data indicate that the body temperature of yearling steers 
are somewhat higher in South Florida than in other parts of the 
country. Average temperature for the Angus, Brahman, and Here- 
ford steers were 104.2, 103.4, and 104.1° F, respectively. Differ- 
ences between breeds were significant. Variations in rectal tem- 
peratures within each breed were significantly correlated with en- 
vironmental temperatures at recording time (0.62). The higher 
rectal temperatures occurred when environmental temperatures 
were the highest. 

Literature Cited 

Bianca, W. 1963. Rectal temperatures and respiratory rate as indicators 
of heat tolerance in cattle. Jour. Agri. Sci., vol. 60, pp. 113-120. 

Bonsma, J. C. 1948. Increasing adaptability by breeding. Farming in South 
Africa, vol. 23, p. 439. 

Brody, S. 1956. Climatic physiology of cattle. Jour. Dairy Sci., vol. 39, 
pp. 715-725. 

Findlay, J. D. 1950. The effects of temperature, humidity, air movement 
and solar radiation on the behavior and physiology of cattle and other 
farm animals. Hannah Dairy Research Institute, Kirkhill, Ayr., Bull., 
no. 9, pp. 19-30. 

Garrett, W. N., T. E. Bond, and C. F. Kelly. 1960. Effect of air velocity 
on gains and physiological adjustments of Hereford steers in a high 
temperature environment. Jour. Animal Sci., vol. 19, pp. 60-66. 

Haines, C. E. 1961. Herefords "run hot" in South Florida. Sunshine State 
Agric. Res. Rept, vol. 6, p. 13. 

Morrison, F. B. 1957. Feeds and feeding. The Morrison Publishing Com- 
pany, Ithaca, New York, ed. 22, pp. 150-155. 

Rhoad, A. O. 1938. Proc. Amer. Soc. Animal Prod., Nov., p. 284. 

Rieck, R. F., and D. H. K. Lee. 1948. Reaction to hot atmosphere of Jersey 
cows in milk. Jour. Dairy Res., vol. 15, p. 219. 



Haines and Kocer: Body Temperatures of Cattle 95 

Snedecor, G. W. 1956. Statistical methods. Iowa State College Press, 
Ames, Iowa, ed. 5. 

Everglades Experiment Station, Belle Glade, Florida and De- 
partment of Animal Science, University of Florida, Gainesville, 
Florida. Florida Agricultural Experiment Stations Journal Series 
No. 1843. 



Quart. Jour. Florida Acad. Sci. 27(1) 1964 



MEDALISTS OF THE FLORIDA ACADEMY OF SCIENCES 

Beginning in 1963, the Florida Academy of Sciences has made 
two annual awards of a medal for basic research to an outstanding 
Florida scientist. Dr. C. C. Clark, of the University of South Flor- 
ida, has served as chairman of the Honors Committee. 

Archie Carr was the medalist for 1963. Born in Mobile, Ala- 
bama, June 16, 1909, Dr. Carr was educated at the University of 
Florida (B.S., 1933; M.S., 1934; Ph.D., 1937). He has since served 
on the staff of that institution, where he is currently Graduate Re- 
search Professor of Biology. Dr. Carr is the author of six books 
and some seventy papers on herpetology and tropical ecology. 
His current research deals with the migration and navigation of 
sea turtles. The recipient of many awards, including membership 
in Phi Beta Kappa, the Elliot medal of the National Academy of 
Sciences, and the Burroughs award, he is a fitting choice for the 
first medal given by the Florida Academy of Sciences. 

The 1964 medalist is Werner A. Baum. Born in Gieszen, Ger- 
many, April 10, 1923, he came to the United States at the age of 
ten. He was educated at the University of Chicago (B.S., 1943; 
M.S., 1944; Ph.D., 1948). After spending two years on the faculty 
of the University of Maryland, he came to Florida State University 
in 1949, where he was successively head of the department of 
meteorology, director of university research, and dean of the grad- 
uate school. In 1963 he was appointed Vice President for Academic 
Affairs and Dean of the Faculties at the University of Miami. Dr. 
Baum has published nearly fifty papers, reviews, and reports. 
Since 1957 he has served as editor-in-chief for the American Mete- 
orological Society and has guided its publications into the category 
of journals of high international stature. Through his own research 
microclimatic investigations have become a significant part of clima- 
tology. The Florida Academy of Sciences is pleased to recognize 
a man who is able to carry on research despite heavy administra- 
tive activities. 



FLORIDA ACADEMY of SCIENCES 

Institutional Members 
for 1964 

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University of Florida 

University of Miami 

University of South Florida 

University of Tampa 



FLORIDA ACADEMY of SCIENCES 

Founded 1936 



OFFICERS FOR 1964 

President: George K. Reid 

Department of Biology, Florida Presbyterian College 

St. Petersburg, Florida 

President Elect: O. E. Frye, Jr. 

Game and Fresh Water Fish Commission 

Tallahassee, Florida 

Secretary: John D. Ktlby 

Department of Biology, University of Florida 

Gainesville, Florida 

Treasurer: John S. Ross 

Department of Physics, Rollins College 

Winter Park, Florida 

Editor: Pierce Brodkorb 

Department of Biology, University of Florida 

Gainesville, Florida 



Membership applications, subscriptions, renewals, changes 

of address, and orders for back numbers should 

be addressed to the Treasurer 



Correspondence regarding exchanges 
should be addressed to 

Gift and Exchange Section, University of Florida Libraries 
Gainesville, Florida 



I * Quarterly Journal 

of the 

Florida Academy of Sciences 

Vol. 27 June, 1964 No. 2 



CONTENTS 

A new crab from the Eocene of Florida Arnold Ross and R. J. Scolaro 97 

White-cedar stands in northern Florida 

E. A. Collins, C. D. Monk, and R. H. Spielman 107 

A new damselfly from the West Indies (Odonata: Protoneuridae) 

Minter J. Westfall, Jr. Ill 



Oxygen depletion in a Florida lake 


George K. Reid 120 


Frogs introduced on islands 


Wilfred T. Neill 127 


Ethoxyquin and vitamin E studies in poultry 

R. H. Harms, C. R. Douglas, 


and P. W. Waldroup 131 


Soil survey for planning urban development 


Victor W. Carlisle 139 


Demography of a Floridian alcoholic sample 


James H. Williams 148 


The shrimp Trachypeneus similis in Tampa Bay 


Carl H. Saloman 160 


Officers and Members of the Academy 


^s^v m 


/ 


^ 


Mailed July 14, 1964 


^^vV*l*» * ff 



Quarterly Journal of the Florida Academy of Sciences 
Editor: Pierce Brodkorb 



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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY of SCIENCES 



Vol. 27 



June, 1964 



No. 2 



A NEW CRAB FROM THE EOCENE OF FLORIDA 

Arnold Ross and R. J. Scolaro 

A carapace of the oxystomatous crab Calappilia was collected 
by the senior author in the summer of 1963 from the Williston 
Formation, Ocala Group, Upper Eocene, near Citra, Florida. A 
second specimen referable to this species was collected a few 
months later from the same horizon in a limerock quarry near 




Fig. 1. Dorsal view of carapace of Calappilia brooksi, new species. 
Holotype, U.S.N.M. No. 648599. Disc-shaped foraminiferans at left are 
Lepidocyclina ocalana Cushman. Actual width of specimen 21.9 mm. 



MfflffiMt 



JUL 2 W64 



98 Quarterly Journal of the Florida Academy of Sciences 

Zuber, Florida. After searching through the Florida State Museum 
collections at the University of Florida two additional specimens 
of this species were discovered. Both of these specimens were 
collected by Dr. Harold K. Brooks. Mr. Henry B. Roberts of the 
U. S. National Museum, who confirmed the writers' generic desig- 
nation, pointed out that the specimens represented a new species, 
and informed us that two specimens of this undescribed species 
were housed in the Wagner Free Institute of Science collections. 
These two specimens were loaned to the authors by the Director 
of the Institute, Robert Chambers, Jr. The seventh specimen re- 
ferable to this species was collected by the junior author from the 
same horizon in a limerock quarry about six miles west of Gaines- 
ville, Florida. 

This is the first reported occurrence of Calappilia in Florida, 
and represents an extension of the known geographical distribu- 
tion. Calappilia is represented in North and Central America by 
two species, C. hondoensis Rathbun (1930), at present known only 
from the Tepetate Formation, Upper Eocene, of Baja California, 
Mexico, and C. diglypta Stenzel (1934), known only from the Crock- 
ett Member of the Cook Mountain Formation, Middle Eocene, of 
Texas. A third, but unidentifiable species of this genus was re- 
ported by Roberts (1956) from the Vincentown Formation, Lower 
Eocene (?), of New Jersey. 

Calappilia bonairensis van Straelen (1933) was described from 
deposits of Upper Eocene Age on the island of Bonaire, Nether- 
lands West Indies. This is the only record of Calappilia in the 
Caribbean or South American regions. 

In Europe Calappilia is represented by six species and one 
variety: C. verrucosa Milne-Edwards (1873) and C. sexdentata 
Milne-Edwards (1876), middle Oligocene of southwestern France; 
C. perlata Noetling (1885), lower Oligocene of Germany; C. incisa 
Bittner (1886), middle Eocene of Italy; C. dacica Bittner (1893), 
upper Eocene of Hungary; C. vicetina Fabiani (1910), upper Olig- 
ocene of Italy; and C. dacica var. lyrata Lorenthey and Beurlen 
(1929), upper Eocene of Hungary. 

In the East Indies Calappilia is represented by two species: 
C. borneoensis van Straelen (1923), middle Eocene of Borneo; and 
C. bohmi Glaessner (1929), upper Eocene of Java (see Bohm, 1922). 

Calappilia is probably restricted, stratigraphically, to deposits 
of Eocene age although four species are purported, according to 



Ross and Scolaro: New Eocene Crab 99 

original references, to occur in strata of Oligocene age in Europe. 
All of the presently known species, other than the four European 
forms, are found in strata ranging from lower to upper Eocene as 
indicated above. There appears to be some question, on the part 
of the writers, that these early European age designations are still 
correct. Further studies on the European fauna would undoubt- 
edly help to clarify this question. 

Institutional abbreviations used in the present paper are as 
follows: Florida State Museum, F.S.M.; U. S. National Museum, 
U.S.N.M.; Wagner Free Institute of Science, W.F.I.S. 

Family Calappidae Dana, 1852 

Subfamily Calappinae Alcock, 1896 

Genus Calappilia Milne-Edwards, 1873 

Type Species. Calappilia verrucosa Milne-Edwards, 1873, Mid- 
dle Oligocene (Rupelian) of Biarritz, southwestern France, by 
monotypy. 

Calappilia brooksi, new species 

Diagnosis. The dorsum is strongly tuberculate and divided 
into three distinct regions by two longitudinal furrows. The pos- 
tero-lateral marginal ornamentation consists of seven more or 
less equidistantly spaced, blunt spines or tubercles decreasing in 
size anteriorly. The superior orbital margins are armed with three 
strong, unequal spines. Posterior to the rostrum and just anterior 
to the juncture of the branchio-cardiac and cervical grooves, the 
gastric region bears an arc of four tubercles, the central pair being 
smallest. The anterior branchial region is ornamented with five 
strong protuberances arranged in a circle surrounding a small tu- 
bercle. Two tubercles on the branchial region and three on the 
hepatic form a second circlet on the dorsum. 

Description. The carapace is subglobose, convex, broader than 
long, tuberculate and divided into three distinct areas by two lon- 
gitudinal furrows. The longitudinal furrows which separate the 
median from the lateral areas are broad and deep. The fronto- 
orbital width is about three-sevenths the greatest width of the car- 
apace. The rostrum projects slightly beyond the level of the outer 
orbital angles; it is poorly preserved on all of the specimens, but 
it appears to have been moderately narrow with two divergent 



100 Quarterly Journal of the Florida Academy of Sciences 

points. The orbits are approximately as wide as they are high 
and slant upward and outwards. The superior orbital margins 
are armed with three strong, nearly erect spines. The outer pair 
are convergent and approximately of equal size. The inner spine 
is about twice as wide as the outer spines, and nearly twice their 
height. The lower margins of the orbits are not preserved. 

The marginal ornamentation consists of somewhat blunt tu- 
bercles. The antero-lateral tubercles are small, equi-distantly 
spaced, directed upwards and gradually increase in size posteri- 
orly. The tubercles on both sides of the carapace, at the widest 
point, are broken on all of the specimens. The postero-lateral 
tubercles abruptly increase in size and become more blunt pos- 
teriorly. The second and third postero-lateral tubercles are mod- 
erately small and broader than long. The fourth is approximately 
one-third as long as it is broad. The fifth tubercle is as broad as 
it is long. The third, fourth, fifth and sixth tubercles are more or 
less equidistantly spaced. The distal end of the sixth tubercle is 
broken on the holotype, but it appears to have been broader than 
long. The seventh tubercle is approximately the same size as the 
second and third tubercles. The posterior margin of the carapace 
bears only two moderately short, somewhat triangular tubercles. 
There is no indication of a large spine or tubercle between these 
two tubercles which may have been either granular or smooth as 
presented in the reconstruction (fig. 2). 

The carapace surface is covered with numerous, high, irregular, 
domal tubercles and granules. The gastric region of the carapace, 
posterior to the rostrum and just anterior to the juncture of the 
branchio-cardiac and cervical grooves, bears an arc of two large 
and two small tubercles; the two smallest being situated between 
the larger ones and all being equidistantly spaced. Immediately 
in front of the two lateral tubercles there are two large granules. 
Posterior to the arcuate pattern of four tubercles there are three 
large tandemly arranged tubercles, the most anterior of which has 
associated with it just anterior and laterally on each side one small 
granule. The most posterior tubercle is situated in the urogastric 
region. The cardiac region bears three tubercles and seven gran- 
ules. The granules surround the most anterior of the three tuber- 
cles. The intestinal tubercle is transversely elongated, and situ- 
ated at each end and anterior to it there is a single granule. 

The branchial region bears one circlet of five large tubercles 



Ross and Scolaro: New Eocene Crab 



101 



surrounding a more or less centrally situated small tubercle. The 
tubercle nearest to the angle formed by the junction of the bran- 
chio-cardiac and cervical grooves is the second largest tubercle 
in the circlet. Anterior to this tubercle there are two small tu- 
bercles that parallel the cervical groove. The tubercle below and 
medial to the above large tubercle is the largest and highest in 
the circlet. Proceeding outward from the first mentioned tubercle 
there are two equidistantly spaced, medium sized tubercles, the 
outermost of which does not belong to the circlet. 

In front of the middle tubercle of the abovementioned row of 
three tubercles there is another medium sized tubercle. A second 
medium-sized tubercle lies obliquely in front and outward from 
this latter tubercle. These two tubercles border the branchio-cerv- 
ical groove and form the posterior border of a second circlet of 
five tubercles of equal size. The three anterior tubercles of this 
circlet lie in the hepatic region of the carapace. The two most pos- 




Fig. 2. Reconstructed dorsal view of Calappilia brooksi, new species. 
Drawing prepared by Miss Brenda Baer. 



102 Quarterly Journal of the Florida Academy of Sciences 

terior of these three also parallel the branchio-cervical groove. The 
central granule lies midway between these two tubercles. 

Posterior to the branchial circlet of tubercles there are four 
medium-sized tubercles. Three of these are tandemly situated 
below and outward from the largest tubercle in the circlet. The 
fourth one is situated laterally from the first one in the row of 
three. 

Bordering the postero-lateral margin of the shell there is a row 
of granules. These are nearly equidistantly spaced and decrease 
in size anteriorly. 

The undersurface of the carapace is partially exposed on sev- 
eral of the specimens. The antero-lateral wall of the carapace is 
nearly vertical. In the region of the lateral extremity of the car- 
apace the underside appears to abruptly change from nearly verti- 
cal to slightly horizontal. The postero-lateral margin is very sharp 
and concavely vaulted in the region of the fourth and fifth mar- 
ginal tuberculations. 

There are no traces of the chelae or walking legs. One eye 
is preserved on the holotype, but not sufficiently to permit descrip- 
tion. 

Remarks. Calappilia brooksi is most closely related to C. hon- 
doensis Rathbun and C. diglypta Stenzel. It may be distinguished 
from the former species by its two circlets of five tubercles and 
tri-spinate superior orbital margins. The orbits of C. hondoensis 
appear to be moderately granulate, and the dorsum has one circlet 
of six large tubercles surrounding a moderately large tubercle. 
The post-rostral region of C. hondoensis has an arc of four tubercles 
and, "in addition several smaller ones" whereas the present new 
species possesses two large and two small tubercles and two gran- 
ules anterior to the outer large tubercles. The cardiac region of 
C. brooksi new species is ornamented with three tubercles and 
seven granules, the latter surrounding the most anterior tubercle. 
Calappilia hondoensis has one elongate median cardiac tubercle 
surrounded by eight regularly placed smaller ones. The present 
new species may be separated from the latter above-mentioned 
species by the greater number of tubercles, ten of which form two 
circlets on each side of the dorsum, the tri-spinate superior orbital 
margins and the lack of a tri-spinate posterior margin. The cardiac 
region of C. diglypta bears only one small tubercle whereas C. 
brooksi new species bears three tubercles and seven granules. The 



Ross and Scolaro: New Eocene Crab 103 

narrow hepatic region of C. diglypta is ornamented with "three 
small granulate tubercles" in contradistinction to this new species 
which is ornamented with three small tubercles arranged in a 
semi-circle surrounding a granule. Four of the branchial tubercu- 
lations of C. diglypta parallel the branchio-cardiac furrows while 
a second group of four medium tubercles are arranged in a Y be- 
tween the aforementioned row and the antero-lateral margin, thus 
serving to distinguish it from C. brooksi new species which has 
the anterior branchial tuberculations forming a circle. 

Two of the specimens in the type series, only one of which is 
figured (fig. 3e), have larger and better developed tubercles on 
the carapace, although the number of large tubercles and their 
position is the same as the remaining specimens. At the present 
time none of the specimens can be differentiated on the basis of 
sex. However, it is the authors belief that these small, well tu- 
berculated specimens may prove to be males when more and bet- 
ter preserved specimens become available for study. 

Measurements of Holotype. Greatest width of carapace 21.9 
mm., length of carapace 18.4 mm., approximate height 8.4 mm. 

Measurements of Paratypes. U.S.N.M. No. 648600, width of 
carapace 26.2 mm., length 19.9 mm. W.F.I. S. No. 17114a, width 
of carapace 16.6 mm., length 15.7 mm. W.F.I.S. No. 17114b, width 
of carapace 22.1 mm., length 18.1 mm. F.S.M. No. 1331, width of 
carapace 20.5 mm., length 15.1 mm. F.S.M. No. 1332, width of 
carapace 13.9 mm., length 11.2 mm. F.S.M. No. 1333, width of 
carapace 15.0 mm., length 12.6 mm. (specimen not here figured). 

Type Locality and Horizon. Limerock quarry in the S.W. V4, 
sec. 35, T. 13 S., R. 22 E., on the south side of a secondary road 
about 1.7 miles east of U. S. Highway 301. This unmarked road 
is approximately 0.9 miles south of the intersection of Florida High- 
way 318 and U. S. Highway 301, in Citra, Marion County, Florida; 
Williston Formation, Ocala Group, Upper Eocene; Arnold Ross 
collector, June, 1963. 

Paratype Localities and Horizon. Limerock quarry on west 
side of U. S. Highway 441, N.W. V 4 , sec. 23, T. 14 S., R. 21 E., just 
south of Zuber, Marion County, Florida; Williston Formation, 
Ocala Group, Upper Eocene; Arnold Ross collector, August, 1963; 
F.S.M. No. 1332. 

Limerock quarry at town of Haile, sec. 13, T. 9 S., R. 17 E., 
Alachua County, Florida; Williston Formation, Ocala Group, Up- 



104 Quarterly Journal of the Florida Academy of Sciences 

per Eocene; Arthur H. Hopkins collector, December, 1952; W.F.I.S. 
Nos. 17114a and 17114b; Harold K. Brooks collector; F.S.M. No. 
1331. 

Limerock quarry about 0.5 miles north of Florida Highway 26 
and approximately 3.3 miles west of the U. S. Interstate Highway 
75 overpass, N.W. %, sec. 35, T. 9 S., R. 18 E., Alachua County, 




Fig. 3. Dorsal views of paratypic specimens of Calappilia brooksi, new 
species. a, W.F.I.S. No. 17114a, actual width of specimen 16.6 mm. b, 
F.S.M. No. 1331, actual width of specimen 20.5 mm. c, U.S.N.M. No. 648600, 
actual width of specimen 26.2 mm. d, W.F.I.S. No. 17114b, actual width of 
specimen 22.1 mm. e, F.S.M. No. 1332, actual width of specimen 13.9 mm. 



Ross and Scolaro: New Eocene Crab 105 

Florida; Williston Formation, Ocala Group, Upper Eocene; Har- 
old K. Brooks collector, 1961, U.S.N.M. No. 648600; R. J. Scolaro 
collector, March, 1964, F.S.M. No. 1333. 

Type Depositories. The holotype and one paratype are de- 
posited in the collections of the U. S. National Museum, catalogue 
numbers 648599 and 648600, respectively. Two paratypes are 
deposited in the Wagner Free Institute of Science collections and 
are numbered 17114a and 17114b, respectively. The remaining 
three paratypes are deposited in the Florida State Museum collec- 
tions at the University of Florida, catalogue numbers 1331, 1332, 
and 1333. 

Etymology. The authors take great pleasure in naming this 
new species in honor of Dr. Harold K. Brooks, Curator of Inverte- 
brate Paleontology, Florida State Museum. 

Acknowledgments 

The authors are indebted to Mr. Henry B. Roberts of the U. S. 
National Museum for his invaluable criticism of the manuscript. 
For permission to borrow specimens from the Wagner Free In- 
stitute of Science collections the authors would like to thank the 
Director, Robert Chambers, Jr. 

Literature Cited 

Bittner, A. 1886. Neue Brachyuren des Eocans von Verona. Sitzungsbr. 
Akad. Wiss., Vienna, vol. 94, no. 1, pp. 44-55, 1 pi. (original reference 
not seen). 

. 1893. Brachyuren aus Tertiarbildungen von Kroatien. Ibid., vol. 

102, no. 1, pp. 10-37, pi. 2 (original reference not seen). 

B6hm, J., in K. Martin. 1922. Die Fossilien von Java, Crustacea. Samml. 
Geol. Reichsmus., Leiden, n.F., vol. 1, no. 2, p. 527, pi. 63 (original 
reference not seen). 

Fabiani, R. 1910. I crostacei terziarii del Vicentino (Illustrazione di alcune 
specie e catalogo generale delle forme finora signalata nella provincia). 
Boll. Mus. Civ. Vicenza, vol. 1, no. 1, p. 4, pi. 1 (original reference not 
seen). 

Glaessner, M. 1929. Fossilium catalogus; Crustacea Decapoda. vol. 1, 
pt. 41, pp. 1-464. 

Lorenthey, E., and K. Beurlen. 1929. Die fossilen Dekapoden der Lander 
der Ungarischen Krone. Geol. Hungarica, Ser. Palaeon., fasc. 3, pp. 
1-420, 16 pis. 



106 Quarterly Journal of the Florida Academy of Sciences 

Milne-Edwards, A. 1873. In R. de Bouille: Paleontologie de Biarritz et de 
quelques autres localites des Basses-Pyrenees. Compte-rendu Trav. 
Congres Sei. France, session 39, vol. 1, pp. 434-435, pi. 4, fig. 3. 

. 1876. In R. de Bouille: Paleontologie de Biarritz et de quelques 

autres localites des Basses-Pyrenees. Bull. Soc. Sci. Lettr. Arts de 
Pau, for 1875-1876 (original reference not seen). 

Noetling, F. 1884. Die Fauna des Samlandischen Tertiars. Abh. Geol. 
Spezial Karte Preussen und d. Thuring Staaten, vol. 6, pt. 3, pp. 1-216. 
pi. 2 (original reference not seen). 

Rathbun, M. J. 1930. Fossil decapod crustaceans from Mexico. Proc. U. S. 
Nat. Mus., vol. 78, art. 8, pp. 1-10, pis. 1-6. 

Roberts, H. B. 1956. Early Tertiary decapod crustaceans from the Vincen- 
town Formation in New Jersey. Bull. Wagner Free Inst. Sci., vol. 31, 
no. 2, pp. 5-12, pi. 2. 

Stenzel, H. B. 1934. Decapod crustaceans from the Middle Eocene of 
Texas. Jour. Paleon., vol. 8, no. 1, pp. 38-56, pis. 6-7. 

Straelen, V. van. 1923. Description de Crustaces decapodes nouveaux des 
terrains Tertiaires de Borneo. Verh. Kon. Akad. Wetensch., Amsterdam, 
vol. 26, pp. 489-492, 3 figs, (original reference not seen). 

. 1933. Sur les Crusteces decapodes de l'Eocene superieur de l'lle 

Bonaire. Bull. Mus. Royal d'Hist. Nat. Belgique, vol. 9, no. 23, pp. 
1-4, 2 figs, (original reference not seen). 

Department of Geology, University of Florida, Gainesville, 
Florida. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



WHITE-CEDAR STANDS IN NORTHERN FLORIDA 
E. A. Collins, C. D. Monk, and R. H. Spielman 

The Atlantic white-cedar (Chamaecyparis thyoides) extends 
from Maine southward into northern Florida and westward into 
Mississippi (Korstian, 1931). In northern Florida the species is one 
of the rarest native trees. Ward (1963) records the two most south- 
ern stands along Cabbage Creek in Putnam County and along 
Juniper Run in the Ocala National Forest in Marion County. 

Cabbage Creek drains an extensive hydric hammock known 
locally as Miller Hammock. The creek then flows southeast about 
four miles where it joins Deep Creek which flows into the Oklawaha 
River. The range of the white-cedar in this area is approximately 
six miles in length. Only a few scattered trees can be seen within 
the eastern end of Miller Hammock, but as one travels east along 
Cabbage Creek, white-cedar increases in importance, until almost 
pure stands are seen near State Road 315. White-cedar begins to 
decrease in importance just east of the point where Gum Creek 
joins Cabbage Creek. Eastward white-cedar occurs only as scat- 
tered individuals. 

The main objective of this study was to determine the composi- 
tion of these southern white-cedar stands. 

Methods 

The sites along Cabbage and Gum Creeks were chosen for 
sampling. In site number 1 white-cedar formed nearly pure stands. 
In site number 2 white-cedar was common, and in number 3 it 
existed as scattered individuals. 

Each stand was sampled by means of the quarter method (Cot- 
ton and Curtis, 1956). In addition to tree data, composite soil 
samples from 0-6, 6-12, 12-18, and 18-24 inch depth were collected. 
The soil samples were oven dried at 105 °C and passed through 
a 2 mm. sieve. From these samples the following determinations 
were made: (1) pH, (2) ppm calcium, (3) ppm magnesium, (4) ppm 
phosphorus, and (5) ppm potassium. 

Results 

A total of 22 species were sampled as trees (> 4 inches d.b.h.) 
or saplings (< 4 inches d.b.h. > 1 inch d.b.h.) from the three 



108 Quarterly Journal of the Florida Academy of Sciences 



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Collins, Monk. Spielman: White-cedar in Florida 109 

stands. Of these, seven occurred in all three stands, seven in two 
stands, and eight in only one stand (Table 1). 

Stand No. 1 was clearly dominated by white-cedar with only 
three other species, sweet bay, redbay, and loblolly pine (Pinus 
taeda), frequent enough to be significant. Dominant saplings were 
wax myrtle (Myrica cerifera), white-cedar, sweet bay, and redbay. 
Saw palmetto was the main shrub species. 

Stand No. 2 in Cabbage Creek swamp was dominated by white- 
cedar. Redbay {Per sera borbonia), red maple (Acer rubrum), ash 
(Fraxinus caroliniana), and several other species were present. The 
dominant saplings were redbay and hazel alder (Alnus rugosa). 
There was a thick shrub layer of greenbrier (Smilax spp.) and saw 
palmetto (Serenoa repens). 

Stand No. 3 was located in the swamp of Gum Creek which 
flows into Cabbage Creek. The hardwoods formed a dense canopy 
which prevented herbs and shrubs from becoming established. 
Very little litter was on the ground. The land was much drier 
then the other stands. Cabbage palm (Sabal palmetto) was the 
dominant species, followed by laurel oak (Quercus laurifolia), black 
gum (Nyssa bioflora), and sweet bay (Magnolia virginiana). The 
dominant saplings were ironwood (Carpinus caroliniana), sweet 
bay, and black gum. 

The organic layer varied from zero to eighteen inches in depth. 
Below the organic layer was coarse sand grading into gravel. 
Charcoal was present in all samples at all levels. The pH was 
highest in the stand with the greatest amount of white-cedar while 
the nutrient content was lowest in the same stand (Table 2). 

TABLE 2 

Soil characteristics of white-cedar stands in Putnam County, Florida 
Averages from composite samples at depth of 6-12 inches 



Stand 


Calcium 


Magnesium 


Potassium 


Phosphorous 


pH 


Number 


in ppm 


in ppm 


in ppm 


in ppm 




1 


791 


115 


12 


2 


7.5 


2 


2144 


404 


38 


7 


6.7 


3 


1289 


175 


15 


3 


6.6 



110 Quarterly Journal of the Florida Academy of Sciences 

Discussion 

Korstian (1931) stated that white-cedar in North Carolina is 
fire-maintained, but the fire must occur when the ground surface 
is under water or the seeds will be destroyed. Fires appear to 
have been fairly common on these Florida stands, but their role in 
succession is not clear. Some old burn scars were seen up to 
twelve feet on the boles. Perhaps past fires have been adequate 
to impede hardwood invasion. 

The white-cedar stands in Putnam County do not occur on 
deep organic deposits as they do in the more northern areas. Little 
(1950) recorded peat 100 feet in depth beneath white-cedar stands 
in New Jersey. In the present study the maximum peat thickness 
was eighteen inches. The pH of the substratum in New Jersey 
ranged from 4.5 to 5.5, while in Florida the range was 6.6 to 7.5. 

Floristically the Florida stands are more diverse. A total of 
22 tree species occurred. Bernard (1963) in a study of lowland for- 
ests in southern New Jersey sampled only six species, red maple, 
white-cedar, sweet bay, black gum, sweetgum (Liquidambar styrac- 
iflua), and American holly (Ilex opaca). Only the holly was not 
encountered in the Florida stands. 

Literature Cited 

Bernard, John M. 1963. Lowland forest of the Cape May formation in 
southern New Jersey. Bull. New Jersey Acad. Sci., vol. 8, pp. 1-12. 

Cottam, G., and J. T. Curtis. 1956. The use of distance measures in 
phytosociological sampling. Ecol., vol. 37, pp. 451-460. 

Korstian, C. F. 1931. Southern white-cedar. U. S. Dept. Agric, Tech. 
Bull., no. 251, pp. 1-76. 

Little, S., Jr. 1950. Ecology and silviculture of white cedar and associ- 
ated hardwoods in New Jersey. Yale Univ. School of Forestry Bull., 
no. 56, pp. 1-103. 

Ward, D. B. 1963. Southeastern limit of Chamaecyparis thyoides. Rhodora, 
vol. 65, pp. 359-363. 

Department of Botany, University of Florida, Gainesville, 
Florida. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



A NEW DAMSELFLY FROM THE WEST INDIES 
(ODONATA: PROTONEURIDAE) 

MlNTER J. WESTFALL, Jr. 

In the Cornell University collection I had seen a series of Pro- 
toneura sent from Puerto Rico by Garcia-Diaz, under date of De- 
cember 19, 1940. Both males and females were green, with no 
indication of the violet and blue described for P. capillaris (Ram- 
bur) and which I observed on a few poorly preserved males from 
Cuba. This made me doubt that the specimens from Puerto Rico 
were of the same species as those from Cuba. Subsequently I was 
able to collect a good series of P. capillaris in Cuba (Westfall, 1964) 
and to obtain in Jamaica in 1960 specimens that agreed with those 
from Puerto Rico. Study in the laboratory confirmed my suspicion 
that two species were involved. 

Rambur (1842) described Protoneura capillaris from a Cuban 
male lacking the head and end of the abdomen. The female re- 
mained unknown until Selys (1886) described it from Cuba. Kolbe 
(1888) listed the species from Puerto Rico on the basis of speci- 
mens collected by Consul Krug. Carpenter (1896), Calvert (1909). 
and Gowdey (1926) reported it from Jamaica. Williamson (1915) 
stated that it was found only in Cuba. Klots (1932) listed it from 
Cuba, the Isle of Pines, Jamaica, Puerto Rico, and Martinique, 
although I do not know the source of the Martinique record. Gar- 
cia-Diaz (1938) reported a single specimen taken in Puerto Rico. 
Whitehouse (1939, 1943) recorded the capture of a mating pair at 
Envy Gut on St. John, Virgin Islands, December 16, 1937, and 
added that neither Wilson in the summer of 1910 nor he from 
November 1941 to May 1942 had been able to find it in Jamaica. 

When I wrote Dr. Thomas W. Donnelly that I intended to de- 
scribe a new species closely related to Protoneura capillaris, he 
sent me a pair which he had taken on St. Thomas and a long series 
from St. John, all of which were of the new species. On his next 
visit to St. John, Dr. Donnelly collected nymphs at the pool where 
he had captured the adults. These nymphs will be described and 
illustrated in a subsequent paper, along with the nymph of Proto- 
neura capillaris, which has been received from Cuba through the 
kindness of Dr. Pastor Alayo D. 



112 Quarterly Journal of the Florida Academy of Sciences 

P. capillaris is apparently restricted to Cuba, and possibly the 
Isle of Pines athough I have not seen the specimens reported from 
the latter island. All literature records of this species from other 
localities in the West Indies, with the possible exception of that 
from Martinique, should probably be referred to the following 
new species. 

Protoneura viridis, new species 

Description of male holotype. Head mostly shining metallic 
green with blue reflections; distal border of labrum narrowly mar- 
gined with orange; anteclypeus, vertical face of frons adjacent to 
clypeus, a rounded spot on each side anterior to antennal base, 
and genae white; back of head black with slight metallic reflections; 
a narrow white stripe bordering compound eyes and continuous 
with white of genae. Prothorax shining metallic green with bluish 
reflections; lateral margin of anterior lobe, median lobe broadly 
adjacent to procoxae, and a very small area on each lateral ex- 
tremity of hind lobe cream-color. Pterothorax with mesepister- 
num entirely shining metallic green with blue reflections; meso- 
stigmal plates with lateral tips yellow; upper half of mesinfraepis- 
ternum and upper part of mesepimeron shining metallic green, 
which color spreads down across first lateral suture to cover about 
posterior fifth of metepisternum; a narrow stripe toward posterior 
end of humeral suture and dorso-posterior corner of mesepimeron 
cream-colored; a black stripe on metepisternum bordering met- 
epimeron, which continues across dorsal margin of metinfraepister- 
num, and in suture dorsal to metacoxa; metepimeron and venter 
of pterothorax cream-colored except for a black median spot be- 
tween metacoxae. Legs mostly pale with black spines; coxae and 
trochanters cream-colored; femora white on external surfaces with 
a dark band toward apex which is faintly visible from external side, 
more so on first femora; black streak on posterior (extensor) sur- 
faces of all femora; tibiae and tarsi cream-colored to brownish, 
darker on flexor surface; joint between femur and tibia blackish. 
Wings narrow, width a little less than one-seventh the length; 
first antenodal space longer than third, and more than twice second 
antenodal space; R s arises at subnodus, M 3 proximal; M 2 arises 
at fifth postnodal in front wings, and at fourth in hind wings; M 3 
ends distal to stigma; Cui ends at termination of crossvein de- 
scending from subnodus. Abdomen very thin and needle-like, but 



Westfall: New West Indian Damselfly 



113 



end segments markedly swollen; segments 1 and 2 dark above 
with slight metallic green reflections, cream-colored on sides; seg- 
ments 3-6 brown to black, anterior segments tending to show me- 
tallic green reflections on dorsum, narrower anterior light rings, 
sides lighter brown, apices darker; segment 7 with a light basal 
ring covering from two-fifths to one-half the length; posterior por- 
tion of segment 7 and segments 8-10 with reddish bronze reflec- 
tions quite noticeable in good light, with margins of 8 and 9, and 
underside of 10 light. Appendages shaped as shown in figures 









Plate 1. Thorax of Protoneura viridis n. sp. and P. capillaris (Rambur). 
Lateral view of male (figs. 1-2). Lateral view of female (figs. 3-4). Mesostig- 
mal plates of female (figs. 5-6). P. viridis (figs. 1, 3, 5); P. capillaris (figs. 
2, 4, 6). 



114 Quarterly Journal of the Florida Academy of Sciences 

7, 9, and 11, a little longer than segment 10; superiors dark above, 
light below, and longer than the light brown inferiors, which are 
darker at extreme base and at tip. 

Description of allotype female. Head and thorax as in male 
but hind lobe of prothorax more noticeably inclined and entirely 
cream-colored except for a small brown median area on the pos- 
terior surface; mesostigmal plates shaped as shown in figure 5, 
their anterior and posterior elevated margins cream-colored, joined 
by a light band across groove between them; external surfaces of 
femora darker than in male. Abdomen colored about as in male, 
but segment 7 entirely dark above; 9 largely light with a dark basal 
ring covering from one-fourth to one-third of segment length and 
extending dorso-laterally to form a triangular projection each side 
to one-half segment length, also with hind margin narrowly dark- 
ened; segment 10 with the dorsally notched posterior margin 
slightly lighter. Appendages conical and shorter than segment 10; 
ovipositor reaching end of segment 10, mostly light, with postero- 
ventral tips black; stylus brown. 

Measurements. Holotype $ : total length including append- 
ages, 36; abdomen, 31; hind wing, 16.2; hind femur, 2.5; longest 
tibial spine on hind leg, 0.43 mm. Allotype 9 : total length, 33; 
abdomen, 27.5; hind wing, 18.2; hind femur, 2.3; longest tibial spine 
on hind leg, 0.43 mm. Paratype $ $ : total length, 34-38; abdo- 
men, 29.3-33; hind wing, 15.5-16.6 mm. Paratype 2 ? : total length, 
32.5-34; abdomen, 27.5-29; hind wing, 18.9-20.1 mm. 

Variations. The coloration with few minor exceptions is re- 
markably uniform in the entire series. The bluish reflections of 
the head and thorax are less noticeable on some specimens taken 
in Puerto Rico in 1940. This is due to the preservation as the 1963 
specimens show the bluish very well. Occasionally vein M 2 arises 
nearer the sixth postnodal in the front wings, and nearer the fifth 
in the hind wings. Cui sometimes ends a little proximal to the 
crossvein descending from the subnodus, as much as the length 
of half a cell in a few front wings. In no specimens studied does 
it end beyond the crossvein, as it does in P. capillaris. 

Etymology. The name of the species is suggested by the ex- 
tensive green markings of the head and thorax in both sexes. 

Specimens examined. Holotype $ (No. 656) and allotype $ 
(No. 657). L'Esperance, St. John, Virgin Islands, June 20, 1961, 
collected by Thomas W. Donnelly. The holotype and allotype 



Westfall: New West Indian Damselfly 



115 



are in the University of Florida collections at Gainesville, Florida. 
Paratypes (44 $ $ , 20 9 9). St. John: L'Esperance, 18 2 2 , 
10 9 9 (Nos. 658-685), June 20, 1961, and 5 $ $ , 3 9 9 (Nos. 686- 
693), August 18, 1963, collected by Thomas W. Donnelly. St. 
Thomas: Turpentine Run at Donoe, 1 $ , 1 9 in copulation (Nos. 
694 and 695), August 28, 1956, collected by Thomas W. Donnelly. 
Jamaica: St. Thomas Parish, about three miles east of Bath, 5 S $ 
(Nos. 696-700), August 15, 1960, collected by M. J. Westfall, Jr. 
and Peter Drummond; Trelawney Parish, along the Martha Brae 
two miles south of Falmouth, 1 $ (No. 701), collected by Thomas 
H. Farr. Dominican Republic: Maymon River, 1 $ (No. 702), 
June 27, 1940, collected by J. G. Needham. Puerto Rico: sup- 







10 





Plate 2. Male abdominal appendages of Protoneura viridis n. sp. and 
P. capillaris (Rambur). Lateral (figs. 7-8), dorsal (figs. 9-10), and ventral (figs. 
11-12) views. P. viridis (figs. 7, 9, 11); P. capillaris (figs. 8, 10, 12). 



116 Quarterly Journal of the Florida Academy of Sciences 

posedly from Rio Piedras, 9 $ $ , 2 2 2 (Nos. 703-713), December 
19, 1940, collected by Julio Garcia-Diaz; Mayagiiez, stream above 
ponds at Agricultural Experiment Station, 4 $ $ , 4 2 2 (Nos. 714- 
721), August 11, 1963, collected by Thomas W. Donnelly. 

In addition I have labeled as this species four specimens in the 
Cornell University collection which have not been designated as 
paratypes because the specimens are in poor condition or the data 
is uncertain. Two are in alcohol, one a complete but broken male 
from San German, Puerto Rico, December 8, 1939, and the other 
a fragmentary female from Mayagiiez, Puerto Rico, November 7, 
1930. One male without appendages is from Mayagiiez, Puerto 
Rico, October 20, 1930, collected by C. U. Salazar. The fourth is 
a headless male with the pin label in ink "Cartagena 12-4-39 P.L.". 
I assume that this might be from Cartagena Lagoon, Puerto Rico, 
and the P.L. could be the initials of the collector. If the date is 
to be interpreted as December 4 it comes close to the dates we 
have. If it should be April 12 it would be considerably earlier than 
the June 20 record. 

The paratypes are presently in the collections of the University 
of Florida, Cornell University, Thomas W. Donnelly, the Kingston 
Museum, and the Academy of Natural Sciences at Philadelphia. 
Some of these will be distributed to other collections. 

Through the courtesy of Dr. Harold J. Grant, Jr., Curator of 
Insects at the Academy of Natural Sciences of Philadelphia, I have 
studied the male specimen listed from Jamaica by Calvert (1909, 
p. 212) as P. capillaris. The data given was "Kingston, by W. J. 
Fox". There were five pin labels and a piece of wing in paper at- 
tached to the pin. None mentioned the collector's name. One 
written label read, "Needham fig" and another, "Penis draw". A 
printed label read, "COLLECTION Ac. Nat. Sci. Phila." while an- 
other read, "PORTLAND, JAMAICA". On the determination label 
in Calvert's handwriting with his printed reference to the 1909 
paper, p. 212, there was the note, "text has locality as 'Kingston' 
erroneously". I assume the correct locality is Portland Parish. It 
is undoubtedly the specimen from which Kennedy (1917) made the 
drawing for his figure 26 of the penis of "Protoneura capillaris 
Ramb., genotype; Portland, Jamaica, in coll. Calvert." The speci- 
men was in poor condition but was easily determined as P. viridis. 
The pin broke in packing it for shipment to me, and I have removed 
the specimen to a transparent envelope with all pin labels attached 



Westfall: New West Indian Damselfiy 117 

to the back of the data card. It has been returned without para- 
type designation because of its poor condition. 

Comparison with P. capillaris. From the specimens of P. 
viridis reported it is evident that the species has a much wider range 
than the known distribution of P. capillaris, which is only Cuba 
and perhaps the Isle of Pines. The color pattern of the thorax in 
the females as shown in the illustrations is remarkably similar but 
is quite different in males. The violet or gun-metal blue of the 
dorsum of P. capillaris is in distinct contrast to the green with blue 
reflections of P. viridis, both in color and in extent of color pattern, 
which covers much more of the mesepimeron in P. viridis. The 
appendages of the males, though superficially alike in lateral view, 
are remarkably different when studied more critically, as shown by 
the figures. The mesostigmal plates of the females are radically 
different in lateral and dorsal views. Except for the fact that few 
good specimens of the Cuban P. capillaris were available for com- 
parison, it seems strange that this new species could have gone 
unnamed for so long. 

Williamson (1915) does not include P. capillaris in his key to 
Protoneura, but on the basis of the character used in couplet b, Cui 
ending against or at the termination of the crossvein descending 
from the subnodus" as opposed to "Cui produced beyond the cross- 
vein descending from the subnodus", P. capillaris and viridis would 
be separated. The new species P. viridis would be grouped with 
calverti Williamson and corculum Calvert, while capillaris would 
come off with cupida Calvert, amatoria Calvert, aurantiaca Selys, 
and cara Calvert. Perhaps P. capillaris and viridis are not so close- 
ly related as their previous confusion would seem to indicate, al- 
though further study of characters other than venational ones might 
yield key characters which would result in a more natural grouping. 

Habits 

My experience in collecting P. viridis is limited to my 1960 
visit to Jamaica, where I collected five males near Bath. The col- 
lecting site was along a small sluggish ditch flowing through open 
fields. The specimens were taken near the road where small trees 
shaded the ditch. They were difficult to detect, and my assistant, 
Peter Drummond, did not notice them. It was late in the after- 
noon and I determined to return on another day to look for more 



118 Quarterly Journal of the Florida Academy of Sciences 

and to attempt to find the nymph. The next day heavy rains turned 
the ditch into a raging torrent, hundreds of feet wide. 

Since Garcia-Diaz reported taking his single specimen in Puerto 
Rico at lights, Whitehouse (1943) suggested that it might be a dusk 
flyer. However, Donnelly (letter of October 2, 1962) found the 
species abundant and ovipositing at midday on St. John. 

Donnelly further states that pairs drifted out of the forest to 
mate and lay eggs in a series of seepage pools at the bottom of a 
sharply defined gut in the hillside. The largest is about 20 feet 
long, 5 feet wide, and perhaps a foot deep. The pools are fed by 
continual seepage from a large fault immediately above the pools, 
which are the only reasonably fresh water in the vicinity. Sunlight 
touches the water for only a few hours each day. The sides of the 
pool are tufa, and the bottom is covered with gravel and a blackish 
mud. The water is choked with fallen leaves, principally mango. 

Nymphs of this species were found clinging to floating mango 
leaves just beneath the surface. Donnelly also found nymphs of 
Perithemis metella (Selys) and Dythemis rufinervis (Burmeister) 
but failed to find nymphs of Macrothemis celeno (Selys), the re- 
maining local species of Odonata. 

Donnelly postulates that the absence of this species from other 
more or less permanent pools on St. Thomas and St. John is a result 
of its being unable to compete with such species as Telebasis 
dominicana (Selys), Enallagma coecum (Hagen), Cannacria herbida 
(Gundlach), and Lepthemis vesiculosa (Fabricius), which were com- 
mon at most other places but absent from the pool on St. John. 

In Jamaica Peter Drummond and I found Protoneura viridis 
associated along the same ditch with adults of Dythemis rufinervis 
(Burmeister), Erythrodiplax fervida (Erichson), E. umbrata (Lin- 
naeus), Micrathyria didyma (Selys), Enallagma coecum (Hagen), 
Ischnura ramburi (Selys), Leptobasis vacillans Hagen and Neoery- 
thr omnia cultellatum (Hagen). We did not collect nymphs at this 
location. 

Acknowledgments 

The work here reported was made possible by a grant from the 
National Science Foundation. All drawings are the work of Esther 
Coogle, former staff artist for the Department of Biology, Univer- 
sity of Florida. Specimens received and help rendered have been 
acknowledged in other parts of this paper. 



Westfall: New West Indian Damselfty 119 

Literature Cited 

Calvert, Philip P. 1909. Contributions to a knowledge of the Odonata of 
the neotropical region, exclusive of Mexico and Central America. Ann. 
Carnegie Mus., vol. 6, no. 1, pp. 73-280, 9 pis. 

Carpenter, George H. 1896. A contribution towards a list of the dragon- 
flies of Jamaica. Jour. Inst. Jamaica, vol. 2, no. 3, pp. 259-263. 

Garcia-Diaz, Julio. 1938. An ecological survey of the fresh water insects 
of Puerto Rico. 1. The Odonata: with new life histories. Jour. Agric, 
Univ. Puerto Rico, vol. 22, no. 1, pp. 43-97. 

Gowdey, C. C. 1926. Catalogus insectorum jamaicensis: Order Odonata. 
Kingston Dept. Agric. Jamaica, Ent. Bull., vol. 4, no. 1, pp. 1-4. 

Kennedy, C. H. 1917. Notes on the penes of Zygoptera (Odonata). No. 
3. The penes of Neoneura and related genera. Ent. News, vol. 28, 
pp. 289-294, 3 pis. 

Klots, Elsie B. 1932. Insects of Porto Rico and the Virgin Islands, Odo- 
nata or dragonflies. Sci. Sur. Porto Rico and Virgin Islands, vol. 14, 
no. 1, pp. 1-107, 7 pis. 

Kolbe, H. J. 1888. Die geographische Verbreitung der Neuroptera und 
Pseudoneuroptera der Antillen, nebst einer Ubersicht uber die von 
Herrn Consul Krug auf Portoriko gesammelten Arten. Archiv Natur- 
gesch., Jahrg. 54, vol. 1, pp. 153-178. 

Rambur, M. P. 1842. Histoire naturelle des insectes Neuropteres. Paris, 
pp. xvii + 534, 12 pis. 

Selys-Longchamps, Edmond de. 1886. Revision du Synopsis des Agrio- 
nines, pt. 1. Mem. Acad. Sci. Belg., vol. 38, pp. iv + 1-233. 

Westfall, Minter, J. Jr. 1964. Notes on the Odonata of Cuba. Quart. 
Jour. Florida Acad. Sci., vol. 27, no. 1, pp. 67-85. 

Whitehouse, Francis C. 1939. Notes on some tropical dragonflies: St. 
John, Virgin Islands, W.I. and Tahiti, French Oceania. Can. Ent., vol. 
71, pp. 199-201. 

. 1943. A guide to the study of dragonflies of Jamaica. Bull. Inst. 

of Jamaica Sci. Ser., no. 3, pp. 1-67. 

Williamson, E. B. 1915. Notes on neotropical dragonflies, or Odonata. 
Proc. U. S. Nat. Mus., vol. 48, pp. 601-638, pi. 38-44. 

Department of Biology and Florida State Museum, University 
of Florida, Gainesville, Florida. 

Quart. Jour. Florida Acad. Sci. 27(2) 1964 



OXYGEN DEPLETION IN A FLORIDA LAKE 
George K. Reid 

For a period of over two weeks in January 1962 an oxygen de- 
pletion occurred in Lake Maggiore, St. Petersburg, Florida. Dur- 
ing this time more than half a hundred tons of dead fish were 
hauled away by sanitation crews. Certain aspects in the recovery 
of the lake were followed regularly until the lake returned to 
normal. Although emphasis was essentially on temperature and 
on oxygen-carbon dioxide relations, some gross observations on 
plankton and other biological features were made. This note pre- 
sents information obtained from field and laboratory data taken 
from 12 January to 3 February 1962. 

Lake Maggiore, named many years ago for its now limnologi- 
cally famous counterpart in Italy, is a brackish, highly eutrophic 
lake occupying a somewhat triangular basin in South St. Peters- 
burg, Florida (salinity: 1 o/oo; chlorides 527.4 mg/1). The lake 
has a surface area of 327 acres (132 hectares) and maximum depth 
is about two meters. The shoreline length is approximately 3.5 
miles (5.63 kilometers) with a shoreline Development Index of 1.44. 
The bottom is sandy but overlain by considerable organic detritus 
of loose texture. A dense phytoplankton content imparts a "pea- 
soup" appearance to the water throughout most of the year. Bor- 
dering the east side of lake is a public park, and residential areas 
are present along the northeast and north shores. The west and 
south shores consist of marsh and palmetto-pine "flatwoods", as yet 
relatively undeveloped. Drainage from the lake is to Tampa Bay 
through a narrow creek at the northeast corner; a spillway with 
mechanical flash boards controls water level and prevents encroach- 
ment of salt water from the bay into the lake. Considerable rec- 
reational pressure in the form of water skiing and boating is ex- 
erted upon Lake Maggiore. 

On 7 January 1962, the first indication of the impending oxygen 
depletion and consequent mass mortality of fishes was observed 
by a city park custodian. On 10 January the local newspaper car- 
ried accounts of the magnitude of the mortality. At 1700 hrs on 
10 January, assisted by Roger Porter, an undergraduate biology 
major at this institution, I launched our boat through hundreds of 
dead and dying fishes (and a very strong stench) to collect water 



Reid: Oxygen Depletion in a Lake 121 

samples and survey the extent of the mortality. Although city 
cleanup crews had been removing carcasses for two days, the east 
(downwind) shore of the lake was littered with thousands of dead 
or gulping fishes, and countless more were seen at the surface of 
the pelagic zone. A local newspaper reported that over 70 tons 
of fish carcasses were removed from 9 to 14 January; the mortality 
continued until 17 January, however. 

Of some six or seven species affected, the brown bullhead 
(Ictalurus nebulosus) constituted fully 90 per cent of the total mass 
killed. Other species included adult mullet (Mugil cephalus), chan- 
nel bass or redfish (Sciaenops ocellata), bluegill (Lepomis macro- 
chirus), threadfin shad (Dorosoma petenense), and the topminnow 
(Gambusia affinis). 

The first measurements of physico-chemical features were made 
at 1730 hrs. on 10 January. The air temperature was about 18 °C 
and the wind westerly at 15-20 mph. Water temperature was uni- 
formly 16.2 °C from surface to bottom at one meter. Dissolved 
oxygen was zero from surface to bottom (determined by Alsterberg 
azide modification of the Winkler method); free carbon dioxide, as 
determined titrimetrically, was 11.0 ppm both at surface and at one 
meter depth. The pH was 7.0. 

Plankton analyses made on 11 January showed a great abun- 
dance of the blue-green alga Anacystis (= Microcystis) sp., but 
no significant quantities of any other algae. Zooplankton in a 
fresh sample of lake water (approximately 7 liters) consisted of less 
than a dozen copepods and their nauplii; these at first appeared 
dead but became active after an hour or so in the laboratory. 

Beginning on 12 January, 1962, air and water temperature, dis- 
solved oxygen, and free carbon dioxide were measured at least 
twice daily (morning and late afternoon) through 30 January, and 
irregularly from that date through 6 February. Water samples 
were taken in a 3-liter Foerst Water Sampler (Kemerrer type) ap- 
proximately 15 meters from shore at depth equal to the length of 
the sampler (50 cm.). The pH was determined occasionally. These 
data are shown in Fig. 1. 

Not until 23 January, or 13 days after the peak of mortality, did 
the dissolved oxygen concentration exceed 2 ppm. It is interesting 
to note that a gradual increase of oxygen was evident from 12 Jan- 
uary to 14 January, but a decrease occurred during the evening of 
the 14th and morning of the 15th. During daylight hours of the 



D.O. - % SATURATION 



TEMP. 



o o o oooooooo oooOoOM^ciaiowj>g)CDoM^(J' 




C0 2 



PPM. 



Reid: Oxygen Depletion in a Lake 123 

12th and 13th the oxygen concentration increased rather uniformly, 
but on the 14th the late afternoon measurement was the same as 
at noon, and on the following day (15th) was much lower. A tenta- 
tive explanation is found in the observation that on the 14th six 
to eight fast boats, some towing water skiers, operated on the lake 
and, in view of the shallow nature of the lake, this activity may 
have increased decomposition of algae and detritus through mix- 
ing; it was thought by local "authorities" that the action of the 
boats would aerate the lake water. At any rate "recovery" appears 
to have been impeded, for a significant build-up of oxygen did not 
appear again until 21 January. 

Throughout the first 5 days of observations the lake retained 
a greenish color and a thick concentration of phytoplankton. About 
15 January, however, the lake became brownish-yellow and re- 
mained so for several days, becoming rather clear by the 21st. 
Qualitative examinations indicated a conspicuous decrease in 
Anacystis from about 18 January through 26 January. On this lat- 
ter date plankton samples contained large numbers of Myxophyceae 
(Anacystis sp.), Chlorophyceae (Closterium sp.), Bacillariophyceae 
(Stauroneis sp.), and an abundance of large holotrichous ciliates. 

A plankton count (Sedgwick-Rafter cell and Whipple ocular mi- 
crometer) of the organisms on 29 January gave results as follows: 

Diatoms: 13.0 x 10 3 /liter 

Scenedesmus: 57.5 x 10 3 /liter 

Merismopedia: 16.5 x 10 3 /liter 

Anacystis: 4.9 x 10 3 /liter 

Unidentified cells: 115.3 x lOVliter 

Some euglenoid flagellates were present, but no microcrustaceans 
or rotifers were observed in the sample strained from 150 liters of 
lake water. Scenedesmus sp., so abundant at this time, had not 
been noted earlier. 

Of biological interest also during this time of stress was the 
behavior of the brown bullhead. As previously indicated, thousands 



Fig. 1. Physico-chemical data for Lake Maggiore, St. Petersburg, Florida, 
12-30 January 1962. Upper curves depict air (dashed line) and water (solid 
line) temperatures and indicate the rapid response of the shallow lake to 
atmospheric temperature fluctuations. Lower curves represent free carbon 
dioxide (C0 2 ) in the water and dissolved oxygen (D. O.) as per cent satura- 
tion at sea level. Midnight is shown by the vertical bars. 



124 Quarterly Journal of the Florida Academy of Sciences 

were killed in the initial period of the catastrophe, and hundreds 
were seen gulping at the surface as late as 12 January. Then for 
two days, during which the dissolved oxygen content increased to 
slightly over one part per million, no bullheads in distress were 
observed. On 14 February (0 2 : 1-1.3 ppm) several tightly grouped 
aggregations of bullheads were seen near shore in water about 0.3 
m deep; a few were gulping at the surface. The following day the 
aggregations were still present and in the late afternoon 2 : 0.1 ppm) 
all individuals showed signs of distress. On 16 January hundreds 
of bullheads were dead and numerous individuals were gulping; 
the oxygen content did not exceed 0.3 ppm during the day. The 
last evidence of mortality was seen on 17 January. Although the 
oxygen content in the morning was 0.0 ppm on 19 and 20 January, 
it did increase to 0.7 and 0.9 ppm in the afternoon of those days, 
respectively. Among other things, these data corroborate the gen- 
erally accepted high tolerance of bullheads to reduced environ- 
mental oxygen content. Furthermore, under environmental con- 
ditions here prevailing, it appears that brown bullheads in the 
size range of 40-80 mm (standard length) generally do not show 
symptoms of distress until dissolved concentration becomes less 
than about one ppm. 

Occurrences such as the one described above always raise 
questions as to the causative agent or agents responsible for 
such massive phytoplankton mortality. In this specific instance 
we cannot, of course, answer with certainty because limnological 
data for the lake prior to the event are not available. Neverthe- 
less, several possible contributing factors acting singly or in con- 
cert are worthy of consideration. 

An algal bloom typically consists of a single species, which has 
become dominant through succession in the phytoplankton com- 
munity. This succession is governed, for the most part, by species 
differences in metabolic activity and cell reproduction in response 
to environmental factors and, indeed, by incompatabilities among 
the various algal species. A popular explanation for succession and 
relative inability of one form to tolerate another is based on one 
species' production and release of substances (perhaps antibiotic 
in nature) that inhibit the growth of other species. The sudden 
mortality of a dominant species may be due to lack of tolerance 
for its own growth-inhibiting substance (Prescott, 1960). This could 
have obtained in the Lake Maggiore situation. 



Reid: Oxygen Depletion 



Lake 



125 



Abrupt changes in environmental factors when the population 
level of a given species is at a critical stage can have deleterious 
effects upon that form. In the case of Lake Maggiore, rather dras- 
tic fluctuations in both air temperature and per cent possible sun- 
shine are found in available climatological data for a period pre- 
ceding the oxygen depletion (Fig. 2). 




I 3 5 



9 II 13 . 15 17 19 21 23 25 27 29 31 
JANUARY- 1962 



3 5 7 9 II 
FEBRUARY 



Fig. 2. Maximum-minimum atmospheric temperatures and prevailing 
sunlight in the vicinity of Lake Maggiore, St. Petersburg, Florida, for the 
period proceding oxygen depletion. Sunshine data were taken by U. S. 
Weather Bureau, Tampa International Airport. During the season under con- 
sideration weather conditions are somewhat widespread and therefore ap- 
plicable to St. Petersburg. 

During the period between 19 and 26 December the minimum 
air temperature dropped nearly 20° C, and from 15 December to 
5 January the general climate underwent a relatively sharp cooling 
followed by rapid warming. Since water temperature in the shal- 
low lake is sharply affected by air temperature (Fig. 1), the atmos- 
pheric variations were obviously reflected in the lake and may have 
contributed to the breakdown of the phytoplankton bloom. 

Especially conspicuous, and probably a significant factor in the 
oxygen depletion, are the three days (29, 31 December, and 6 Jan- 
uary) when less than 10 per cent of possible sunshine prevailed 
(Fig. 2). If the high concentration of algae were conditioned by 



126 Quarterly Journal of the Florida Academy of Sciences 

a given average level of light intensity and duration, this marked 
diminution may well have been sufficient stimulus for the catas- 
trophe. 

Acknowledgment 

This investigation was made in conjunction with a more com- 
prehensive study of Florida lakes supported by National Science 
Foundation grant number 17865. 

Literature Cited 

Prescott, G. W. 1960. Biological disturbances resulting from algal popu- 
lations in standing waters. In Tryon, C. A. Jr. and R. T. Hartmann 
(ed.). The ecology of algae. Spec. Publ. No. 2, Pymatuning Labora- 
tory of Field Biology, University of Pittsburg. 

Florida Presbyterian College, St. Petersburg, Florida. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



FROGS INTRODUCED ON ISLANDS 

Wilfred T. Neill 

The specimens described below are in the herpetological col- 
lection of the University of Florida. I am indebted to Dr. Walter 
Auffenberg for permission to examine and describe them. Thanks 
are also extended to Dr. J. C. Dickinson, Jr., Mr. Thomas G. Baker, 
and Dr. Carter R. Gilbert, for useful information. 

1. Rana grylio on New Providence Island 

The natural range of the pig frog, Rana grylio Stejneger, ex- 
tends from Florida northward into South Carolina and westward 
into southeastern Texas. 

Four young adults of Rana grylio (UF 17209-12) and one tad- 
pole (UF 17213) were collected from Lake Killarney, New Provi- 
dence Island, Bahamas, by Dr. Dickinson and Martin Dickinson, 
on August 12, 1958. The adult frogs include one male, 74 mm. in 
snout-vent length, and three females ranging in snout-vent length 
from 59 to 67 mm. The tadpole is 63 mm. in total length. Addi- 
tional tadpoles (UF 17512-14) were taken in the lake by Dr. Dick- 
inson on August 15, 1960. Evidently the species is breeding at 
the locality. 

Dr. Dickinson was informed that the original stock had been 
deliberately introduced a few years previously, and had been ob- 
tained from a commercial source in Florida. The Bahaman speci- 
mens agreed very closely with those from central Florida. 

Dr. Dickinson, who collected fishes in Lake Killarney in five 
different years, was impressed by the relative scarcity of small 
organisms, of kinds that might provide food for Rana grylio. The 
fish fauna, as identified by Dr. Gilbert, included Cyprinodon varie- 
gatus, Gambusia manni, Rivulus cf. marmoratus, Eleotris amblyop- 
sis, and Mugil sp. Damselflies, dragonflies, and aquatic hemipte- 
rans were also noted. The lake is weakly saline, reminiscent of 
tidal flats in general aspect. It would not seem to provide an opti- 
mum habitat for the pig frog, although this species has occasionally 
been found about brackish marshes in the United States (Neill, 
1958, p. 12). 



128 Quarterly Journal of the Florida Academy of Sciences 

2. Rana catesbeiana on Cuba 

The bullfrog, Rana catesbeiana Shaw, occur over much of east- 
ern North America, and has also been introduced into many areas 
outside its natural range. Its presence on Cuba has been noted 
previously (Stejneger and Barbour, 1939, p. 44), but specific rec- 
ords for the island are lacking, and little is known about the abun- 
dance of the species there. 

Three specimens (UF 17471-73) were collected by me at Rancho 
Mundito, at that time the estate of Col. Fulgencio Batista, in low 
mountains overlooking Consolacion del Sur, Pinar del Rio Province, 
Cuba. Snout-vent lengths ranged from 41 to 75 mm. A few 
larger examples were also seen here. On the Rio Hondo, in the 
lowlands just west of Consolacion del Sur, a young adult bullfrog 
was disgorged by a snake, Tretanorhinus variabilis, and a second 
young adult (WTN 2845) was collected from beside the stream. 
The latter frog measured 52 mm. in snout- vent length. Yet an- 
other small bullfrog (WTN 2846), 48 mm. in snout- vent length, was 
collected between Giiines and Playa de Rosario, Habana Province. 
All the above-mentioned Cuban examples were taken in October, 
1949. 

Bullfrogs from Florida and the lower Coastal Plain of the south- 
eastern United States are heavily dark-mottled above and below; 
specimens from above the Fall Line are usually plain greenish 
above and whitish below. Cuban examples resemble the latter 
population, and not the Florida one. 

The bullfrog was more abundant on the Batista estate than at 
any other Cuban locality investigated; but even at Rancho Mundito 
it was by no means as common as in many parts of its natural 
range. 

Cuba has no native frog similar to the bullfrog in habits. The 
giant Cuban toad, Bufo peltacephalus (Tschudi), roughly equals the 
bullfrog in size, but is often found in dry situations where the 
Rana is lacking. The Bnfo also frequents stream and lake margins, 
where it might compete with the bullfrog. On the Batista estate 
the toad was far more common than the bullfrog. The next largest 
native frog of Cuba, Hyla septentrionalis Dumeril and Bibron, is 
primarily arboreal and so not a frequent competitor of the bull- 
frog. However, during and immediately after hard rains, the 
Hyla will descend to forage on the ground. This treefrog was ex- 



Neill: Frogs introduced on Islands 129 

tremely abundant on the Batista estate. Thus at Rancho Mundito 
the two large native frogs did not appear to suffer from competi- 
tion with the introduced species. 

3. Hyla aurea aurea on New Caledonia 

The green-and-gold bellfrog, Hyla aurea aurea (Lesson), is 
native to eastern New South Wales, Australia. Moore (1961, p. 
316) mentioned three examples from New Caledonia. 

Seven individuals of Hyla aurea aurea (UF 17327-33) were taken 
near Noumea, New Caledonia, by Mr. Baker on June 5, 1962. The 
frogs were found on the road at night, in a low spot near a lake 
on the southern outskirts of the town. 

In this lot the snout-vent length ranged from 58 to 71 mm. The 
dorsum was smooth; the dorsolateral fold, while low, was continu- 
ous, and was conspicuous by virtue of its pale coloration. A mid- 
dorsal stripe was lacking. All the specimens were referable to 
Hyla aurea aurea, not H. aurea raniformis (Keferstein). The lat- 
ter subspecies may also have been introduced into New Caledonia, 
where one specimen has been taken (Moore, loc. cit.). 

New Caledonia, lying in the South Pacific about 800 miles east 
of Australia, harbors no native frogs. Hyla aurea aurea probably 
arrived as a stowaway on ships, there being several large seaports 
within its natural range. 

4. Xenopus laevis on Ascension Island 

The African clawed frog, Xenopus laevis laevis (Daudin), is 
native to southern Africa. 

Three specimens (UF 16000-02) are from Ascension Island, col- 
lected at Bates' Tank on July 9, 1958, by Richard G. Allan. In 
all of this lot the so-called "tentacle" was minute. The habitus 
was pyriform. The ratio of body width to head width fell be- 
tween 1.4 and 1.5. The black claws, viewed from above, appeared 
flattened. The venter was immaculate, although the under surfaces 
of the hind limbs were weakly speckled in the two larger individu- 
als. Snout-vent length ranged from 54 to 66 mm. The frogs 
are probably young adults of the subspecies Xenopus laevis laevis, 
as defined by Loveridge (1933, pp. 351-352). 

Ascension Island, lying in the southern Atlantic about midway 
between Africa and South America, harbors no native frogs. The 



130 Quarterly Journal of the Florida Academy of Sciences 

origin of its Xenopus colony is not known. However, during the 
1930's and 1940's, X. laevis was thought to be exceptionally useful 
in the early diagnosis of human pregnancy. Living specimens came 
to command a high price, and many colonies of the frog were vir- 
tually exterminated in parts of southern Africa. Efforts were made 
to breed the species in captivity, but with little success (Rose, 1950, 
pp. 16, 30). This circumstance may well have led to the attempted 
establishment of new colonies at points outside the natural range. 

Literature Cited 

Loveridge, Arthur. 1933. Reports on the scientific results of an expedition 
to the southwestern highlands of Tanganyika Territory. VII. Herpe- 
tology. Bull. Mus. Comp. Zool., vol. 74, pp. 197-416, pis. 1-3. 

Moore, John A. 1961. The frogs of eastern New South Wales. Bull. Amer. 
Mus. Nat. Hist., no. 121, pp. 149-386, pis. 27-46. 

Neill, Wilfred T. 1958. The occurrence of amphibians and reptiles in 
saltwater areas, and a bibliography. Bull. Marine Sci. Gulf and 
Caribbean, vol. 8, pp. 1-97. 

Rose, Walter. 1950. The reptiles and amphibians of southern Africa. 
Maskew Miller, Ltd., Cape Town, xxv and 378 pp. 

Stejneger, Leonhard, and Thomas Barbour. 1939. A check list of North 
American amphibians and reptiles. 4th ed. Harvard Univ. Press, Cam- 
bridge, Mass., xvi and 207 pp. 

Florida State Museum, Gainesville, Florida. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



ETHOXYQUIN AND VITAMIN E STUDIES IN POULTRY 
R. H. Harms, C. R. Douglas, and P. W. Waldroup 

The addition of 0.0125 per cent of ethoxyquin to diets has been 
reported to increase body weights and increase deposition of pig- 
ment in the skin and shanks of broilers (Waldroup et ah, 1960). 

Machlin et at. (1959) reported that ethoxyquin would prevent 
vitamin E deficiencies in chickens. These workers also reported 
that ethoxyquin or vitamin E was effective in preventing liver 
"peroxide" formation and suggested that this might explain the 
reports by Hickman et al. (1944) and Sherman (1941) that vitamin 
A is destroyed more rapidly in vitamin E deficient rats. 

The experiments reported herein were conducted to determine 
whether the growth promoting activity of ethoxyquin was related 
to the in vivo protection of vitamin A. 

Experiments 1 and 2: Chick Studies, Practical Diets 

Procedure. Four-hundred day-old broiler type chicks, which 
had been intraocularly vaccinated for infectious bronchitis and 
Newcastle disease, were used in each of these experiments. They 
were randomly assorted into forty groups of five males and five 
females each, and placed into electrically heated battery brood- 
ers with raised wire floors. 

A 2 x 2 x 2 factorial arrangement of treatments was used in 
each experiment. This included zero or 10 I.U. of vitamin E per 
pound of diet, supplied as d alpha tocopherol acetate, zero or 
0.0125 per cent of supplemental ethoxyquin, and zero or 3000 I.U. 
of vitamin A per pound of diet. Rasal diet 1 (Table 1) was used 
in this experiment. 

Five pens of chicks were placed on each of eight diets at one- 
day of age. Experimental diets and tap water were given ad 
libitum throughout the experiment. All chicks receiving supple- 
mental vitamin A in the diet were individually weighed at four 
weeks of age, when this phase of each experiment was terminated. 
The chicks receiving no supplemental vitamin A were maintained 
on their respective experimental diets until all had died from a 
vitamin A deficiency. 



132 Quarterly Journal of the Florida Academy of Sciences 



TABLE 1 

Composition of basal diets (lbs/cwt) 









Diet Number 




Ingredients 


1 


2 


3 


4 


5 


Degerminated corn 


65.2 











52.5 


Soybean meal (50% protein) 


31.0 


— 


— 


— 


41.0 


Assay protein C-l 1 


— 


35.0 


35.0 


35.0 


— 


Cerelose 


— 


45.9 


55.9 


49.9 


— 


Stabilized animal fat 2 


— 


— 


— 


6.0 


— 


Ethyl lineoleate 


— 


10.0 


— 


— 


— 


Ground limestone 


0.7 


— 


— 


— 


1.5 


Defluorinated phosphate 












(34% Ca & 18% P) 


1.8 


— 


— 


— 


3.9 


Iodized salt 


0.4 


— 


— 


— 


0.4 


Micro-ingredients 3 


0.9 


— 


— 


— 


— 


Glycine 


— 


1.0 


1.0 


1.0 


— 


Choline CI. (25%) 


— 


0.8 


0.8 


0.8 


— 


Methionine hydroxy analogue (90%) 


— 


0.7 


0.7 


0.7 


0.05 


Vitamin A (10,000 I.U/gm.) 


— 


2 gm 


2 gm 


— 


— 


Vitamin D 3 (3,000 I.C.U./gm.) 


— 


0.01 


0.01 


0.01 


— 


Aureomycin (10 mg./lb.) 


— 


0.1 


0.1 


0.1 


— 


Salts 4 


— 


6.0 


6.0 


6.0 


— 


Vitamin mix 1 


— 


0.5 


0.5 


0.5 


— 


Ethoxyquin 


— 


0.01: 


15 0.0125 


— 


— 


Micro-ingredients 5 


— 


— 


— 


— 


.65 



1 An isolated soybean protein obtained from Archer-Daniels-Midland Co., 
Cincinnati, Ohio. 

2 Stabilized with B.H.T. and B.H.A. 

3 Supplied per pound of diet: 340 I.C.U. vitamin D 3 , 10 meg. vitamin Bi 2 , 
2 mg. riboflavin, 9 mg. calcium pantothenate, 18 mg. niacin, 261 mg. choline 
cl., 10 mg. terramycin, 35 mg. manganous oxide, 9 mg. iron, 0.9 mg. copper, 
90 meg. cobalt, 5 mg. iodine, 45 meg. zinc, and 80 mg. manganese sulphate. 

* According to Machlin and Gordon (1959). 

5 Supplied per pound of diet: 600 I.C.U. vitamin D 3 , 6.3 meg. vitamin 
B12, 2 mg. riboflavin, 9.6 mg. calcium pantothenate, 10 mg. niacin, 120 mg. 
manganese sulfate, 400 mg. choline cl., 10 mg. terramycin, 25 mg. nitrofura- 
zone, and 3.6 mg. furazolidone. 

Statements of probability in this paper are based on the anal- 
ysis of variance according to Snedecor (1956), with significant dif- 
ferences between treatment means determined by Duncan's mul- 
tiple range test (1955). 

Results. In experiment 1, supplementing the diet with 0.0125 
per cent of ethoxyquin in the presence of 3000 I.U. of vitamin A 



Harms et al: Ethoxyquin and Vitamin E 133 

per pound resulted in significantly increased body weight of chicks 
at four weeks of age (Table 2). However, in experiment 2, body 
weights of chicks receiving supplemental ethoxyquin were numeri- 
cally but not significantly lower than those receiving the unsup- 
plemental diets. The observation that ethoxyquin improved growth 
rate in experiment 1 would agree with the findings of Waldroup 
et al. (1960). Supplementing the diet with 10 I.U. of vitamin E per 
pound of diet resulted in a numerically improved growth rate in 
each experiment (Table 2). This improvement in growth was 
statistically significant in experiment 2. It was rather surprising 
that vitamin E improved growth of broilers, although the degerm- 
mated corn meal used in the basal diet resulted in a relatively low 
level of vitamin E. However, this was not repeated in subsequent 
experiments. Since the vitamin E and ethoxyquin interaction was 
not significant it would appear that the growth promoting effect 
of ethoxyquin was not due to a sparing effect on vitamin E. 

TABLE 2 

Body weight and survival time of chicks as affected by Supplemental 
ethoxyquin or vitamin E (Experiments 1 and 2) 

Supplement 4- Week Body Weight 

Ethoxy- (gms) 1 - 2 Survival time days V 

Vitamin E 4 quin 5 Exp 1 Exp 2 Av. Exp 1 Exp 2 Av. 



+ 



Av. 



— 


370a 


425b 


398 


21.2a 


30.0 


25.6 


+ 


389b 


422b 


406 


21.4a 


31.0 


26.2 


Av. 


380 


424 


402 


21.3 


30.5 


25.9 


— 


374a 


446a 


410 


24.9b 


30.6 


27.8 


+ 


394b 


435ab 


415 


23.2ab 


31.3 


27.6 


Av. 


384 


441 


413 


24.1 


31.0 


27.7 


— 


372 


436 


404 


23.1 


30,3 


26.7 


+ 


392 


429 


412 


22.8 


31.2 


26.9 



1 Means not having common exponential letters are significantly different 
according to Duncan's multiple range test (1955). 

2 These chicks received diets containing 3000 I.U. vitamin A per pound. 

3 These chicks received basal diet 1 containing no supplemental vitamin A. 

4 Indicates 10 I.U. supplemental vitamin E/pound. 

5 Indicates 0.0125 per cent supplemental ethoxyquin. 

Survival time of chicks was prolonged by the addition of 10 
I.U. of vitamin E per pound of diet (Table 2). This difference was 



134 Quarterly Journal of the Florida Academy of Sciences 

statistically significant in experiment 1 but not significant in experi- 
ment 2. This finding would agree with results reported by Hick- 
man et al. (1944) who found that survival time of rats after vita- 
min A supplementation ceased was increased by feeding vitamin 
E. The addition of ethoxyquin to the diet numerically increased 
survival time in experiment 2, but not in experiment 1 (Table 2). 
Although the supplemental vitamin E significantly increased sur- 
vival time in experiment 1, these data would suggest that neither 
E nor ethoxyquin was effective in protection of vitamin A in day- 
old chicks. 

Experiment 3: Chick Study, Purified Diet 

Procedure. Six hundred day-old broiler type chicks, which 
had been intraocularly vaccinated for infectious bronchitis and 
Newcastle disease, were randomly assorted into sixty groups of 
five males and five females each, and placed into electrically heated 
battery brooders with raised wire floors. 

One half of the groups were placed on basal diet 2, and the 
other half on basal diet 3 (Table 1). Ethyl lineoleate was included 
in one diet in an effort to increase the level of unsaturated fatty 
acids which would undergo peroxidation in the liver. This ma- 
terial was used since Machlin et al. (1959) suggested that peroxida- 
tion might be a factor in the rate of vitamin A destruction in the 
liver. 

At seven days of age one-third of the groups previously fed 
each of diets 2 and 3 were changed to basal diet 4 (Table 1), an- 
other one-third were changed to basal diet 4 supplemented with 
0.0125 per cent of ethoxyquin, and the remaining one-third re- 
ceived basal diet 4 supplemented with 10 I.U. vitamin E per pound 
of diet. All birds were individually weighed at four weeks of age. 
At this time no visible vitamin A deficiency symptoms had been 
observed. All chicks received their respective diets until all had 
died. 

Results. The addition of ethoxyquin to the diet resulted in 
numerically increased body weight at four weeks of age (Table 
3). This difference in body weight was not affected by the diet 
which the chicks received during the first seven days of life. This 
would indicate that the growth response from ethoxyquin was not 
dependent upon the protection of unsaturated fatty acids from 



Harms et al: Ethoxyquin and Vitamin E 



135 



peroxidation. The addition of vitamin E to the basal diet resulted 
in a slight numerical reduction in body weight. No interaction 
was observed between vitamin E and ethoxyquin, again indicat- 
ing that the response from ethoxyquin is independent of the vita- 
min E level in the diet. 

TABLE 3 

Body weight and survival time of chicks as affected by composition of diet 
or supplemental ethoxyquin or vitamin E 

(Experiment 3) 



0-7 da 


Diet 

7-70 da 


Supple- 
mental 
7-70 da 


4- Week Body Weight (gms) 
M F Av 


Survival 
Time 

(Days) 


Basal 2 


Basal 4 


None 


466 


439 


452 


56,3 






+ S 1 


475 


447 


461 


54.8 






+E 2 


464 


412 


438 


55.6 






All 


468 


433 


450 


55.6 


Basal 3 


Basal 4 


None 


478 


414 


446 


57.6 






+ S 


476 


429 


453 


53.4 






+ E 


486 


405 


446 


55.0 






All 


480 


416 


448 


55,3 


Av. 


Basal 4 


None 


472 


426 


449 


57.0 






+s 


476 


438 


457 


54.1 






+ E 


475 


409 


442 


55,3 



1 Indicates 0.0125 per cent supplemental ethoxyquin. 

2 Indicates 10 I.U. supplemental E/pound. 

The addition of ethoxyquin or vitamin E to the diet resulted 
in numerically reduced survival time of chicks (Table 3). This 
again indicates that neither the antioxidant nor vitamin E was 
effective in protecting vitamin A as measured by survival time on a 
vitamin A deficient diet. However, it is possible that the slight 
accelerated growth resulting from ethoxyquin supplementation may 
have increased the vitamin A needs; thus survival time would not 
have been a true indication of vitamin A protection. 



Experiment 4: Turkey Poult Study 

Procedure. Three-week-old male Beltsville White poults which 
had been maintained on a commercial turkey starter feed were 



136 Quarterly Journal of the Florida Academy of Sciences 

used in this experiment. They were individually weighed and 29 
poults were placed at random into five, 10 x 12 foot range houses 
with wood shavings used as litter. 

Basal diet 5 (Table 1) was used in this experiment. Two groups 
of poults received the basal diet, two groups received the basal 
diet with 0.0125 per cent supplemental ethoxyquin, and one group 
received the basal diet supplemented with ethoxyquin and 3000 
I.U. of vitamin A per pound of diet. The latter diet served as 
the positive control. 

Individual body weights were obtained at weekly intervals un- 
til the poults had been on experiment four weeks. Records were 
kept of the date on which each bird died from vitamin A deficiency. 
The positive control group was terminated after four weeks with 
the remaining poults receiving their respective diets until all had 
died. 

Results. Poults receiving the basal diet supplemented with 
ethoxyquin grew more rapidly during the second and third week 
than did those receiving the basal diet (Table 4). This difference 
in growth rate was significant during the third week. During the 
fourth week the poults began exhibiting vitamin A deficiency symp- 
toms, and those receiving the supplemental ethoxyquin grew more 
slowly than those receiving the basal diet. It is possible that this 
change in growth pattern may have been due to the faster rate of 
growth during the first three weeks, with depletion of vitamin A 
storage at a more rapid rate than in the controls. 

TABLE 4 

Body weight gains and survival time of turkey poults 
fed diets with and without ethoxyquin 

(Experiment 4) 



Diet 










Survival 






Weekly 


gains (gms) 




time 




1 


2 


3 


4 


(days) 


Basal 


177 


207 


287 


92 


33.8 


+0.0125% ethoxyquin 


166 


217 


311* 


84 


33.2 


+ 0.0125% ethoxyquin and 


179 


217 


328 


318 





3000 I.U. vitamin A/lb. 













Significantly different from basal. 



Harms et al: Ethoxyquin and Vitamin E 137 

No significant difference was detected in survival time between 
the group receiving the basal diet and those receiving the diet 
containing the supplemental ethoxyquin (Table 4). This would 
indicate that either the ethoxyquin did not afford any protection 
for vitamin A in turkey poults, or their accelerated growth caused 
a greater demand for the vitamin. 

Summary and Conclusions 

Four experiments were conducted to determine the effect of 
supplemental ethoxyquin and/or vitamin E upon growth rate and 
protection of vitamin A in chicks and turkey poults. 

The addition of 0.0125 per cent ethoxyquin to the diet resulted 
in increased body weights in three of four experiments. The ad- 
dition of vitamin E to the diet increased body weight in one experi- 
ment and decreased it in another, with the average of three exper- 
iments indicating no response. The vitamin E and ethoxyquin in- 
teraction was non-significant, indicating that the growth response 
from ethoxyquin was independent of the vitamin E level of the 
diet. 

The addition of vitamin E to the diet in experiment 1 signifi- 
cantly prolonged life when chicks were placed on a vitamin A de- 
ficient diet, although the average of survival time of chicks in all 
experiments indicated that neither vitamin E or ethoxyquin was 
protecting the vitamin A. However, increased growth resulting 
from ethoxyquin supplementation may have increased vitamin 
needs, and thus survival time might not be a true indication of 
vitamin A protection. 

Acknowledgment 

This investigation was supported in part by a grant-in-aid from 
Monsanto Chemical Company, St. Louis, Missouri. 

Literature Cited 

Duncan, D. B. 1955. Multiple range and multiple F tests. Biometrics, vol. 
11, pp. 1-42. 

Hickman, K. C, M. W. Kaley, and P. L. Harris. 1944. Covitamin studies. 
1. The sparing action of natural tocopherol concentration on vitamin 
A. Jour. Biol. Chem., vol. 152, p. 303. 



138 Quarterly Journal of the Florida Academy of Sciences 

Machlin, L. J., R. S. Gordon, and K. H. Meisky. 1959. The effect of anti- 
oxidants on vitamin E deficiency symptoms and production of liver 
"peroxide" in the chicken. Jour. Nutrition, vol. 67, pp. 333-343. 

Sherman, W. C. 1941. Activity of alpha tocopherol in preventing antag- 
onism between linolenic esters and carotene. Proc. Soc. Exp. Biol. 
Med., vol. 47, p. 199. 

Snedecor, G. W. 1956. Statistical methods. Iowa State College Press, 
Ames, Iowa, ed. 5, 534 pp. 

Waldroup, P. W., C. R. Douclas, J. T. McCall, and R. H. Harms. 1960. 
The effects of santoquin on the performance of broilers. Poultry Sci., 
vol. 39, pp. 1313-1317. 

Department of Poultry Science, University of Florida, Gaines- 
ville, Florida. Florida Agricultural Experiment Stations Journal 
Series No. 1711. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



SOIL SURVEY FOR PLANNING URBAN DEVELOPMENT 

Victor W. Carlisle 

The fundamental purpose of the soil survey, like that of other 
research, is to obtain information necessary to make predictions 
(Kellogg, 1955). Soil surveys are made to accurately record the 
physical characteristics of an area by presenting a natural classifi- 
cation of soils based on their inherent qualities. With organized 
knowledge of soil characteristics, it is possible to make accurate 
predictions of soil behavior. The soil map and our knowledge of 
how soils respond have been used successfully for many years as 
a basis for the most intelligent land use by our rural population. It 
also provides the best basis for planning communities, including 
those in the urban fringe. 

Historical 

Although mapping of soils in the United States began in 1899, 
it has been only in recent years that planners and developers are 
becoming increasingly aware of the wealth of information con- 
tained in soil surveys that may be used for designing a unified 
plan for land use. Use of detailed soil surveys is absolutely nec- 
essary in urban areas if those responsible for the use of the land 
are to make decisions based on facts. 

Fairfax County, Virginia, was one of the first in the country to 
use soil maps for urban planning. In Fairfax County, the County 
Planning and Zoning Commission, Health Department, Department 
of Public Works, County Department of Schools, and Department 
of Assessments are all using the detailed soil survey (Clayton et al., 
1959; Obenshain et al., 1962). In addition, the county employs a 
soil scientist to interpret the maps for various county agencies and 
advise individual landowners. 

Other areas where soil surveys are being made or have been 
completed for use in urban planning include: Lake County, Illinois; 
Montgomery County, Pennsylvania; Santa Clara County, Califor- 
nia; James Island, South Carolina; San Antonio, Texas; Cincinnati, 
Ohio; Hanover, Massachusetts and 39 other communities embrac- 
ing 640,000 acres; Hartford, Danbury, and Stamford, Connecticut; 
and a 6-county area in Wisconsin administered by the Southeastern 
Wisconsin Regional Planning Commission. Soil maps are also be- 



140 Quarterly Journal of the Florida Academy of Sciences 

ing used for urban planning in New Jersey, Michigan, Missouri, 
and Oklahoma (Marshall, 1963). Planners in the Duval, Dade, and 
the 7-county Cape Kennedy area in Florida need soil survey infor- 
mation for locating suitable residential developments, light indus- 
trial sites, wildlife sites, camp sites, picnic sites, and other recre- 
ational facilities. 

Generalized and Detailed Maps 

Soil surveys may be used at several different levels of general- 
ization for urban planning. Small scale maps, such as a state soil 
map or soil association maps of individual counties, may be used for 
primary planning. These generalized soil maps are helpful in 
portraying the fairly well-defined patterns of geographically associ- 
ated soils; comparing the distribution of soils; and in locating rel- 
atively large areas of soils suitable for some particular use. The 
units of a generalized soil map represent associations of individual 
soils. Generalized soil maps provide the urban-fringe planner 
with a view of the soil resources in an entire area and contribute to 
planning on the county level. 

As planning progresses the areas being considered decrease in 
size and recommendations become more specifice. This creates a 
need for more precise soil information. At this stage, generalized 
soil maps showing geographically associated soils cannot be used 
for it is necessary to have far more detailed soil information. In- 
dividual soils with defined characteristics must be delineated. This 
is done by a systematic examination of the soils in the field and 
laboratory. 

Procedures used in making soil surveys have been described 
in detail (Klingebiel, 1963; Soil Survey Staff, 1951). Soil scientists 
walk over the land, identifying different soils by digging holes and 
examining the soil layers. Boundaries are plotted of areas occurring 
along the landscape that are similar in number, thickness, and rela- 
tive arrangement of layers; proportion of sand, silt, and clay; color; 
chemical composition; shape, size, and consistence of natural ag- 
gragates; and kind of parent material. The slope and any abnor- 
mal removal of soil materials by wind or water are recorded. Soils 
with like properties and characteristics are classified under a com- 
mon name. All of the intricate boundaries of a soil are accurately 
plotted throughout their course using aerial photographs as base 
maps. These detailed soil maps are necessary for comprehensive 
urban planning. 



Carlisle: Soils Survey for Planning Urban Development 141 

Criteria for Evaluating Soils 

Soils delineated on the detailed soil map may now be grouped 
according to their inherent properties or characteristics. Any prop- 
erty deemed important in determining the best use of a given soil 
in urban planning may be evaluated in this manner (Bartelli, 1962). 
Criteria commmonly considered in evaluating soils for urban use 
are as follows: topography, permeability, wetness, depth to bed- 
rock, shrink-well potential, bearing values, susceptibility to erosion, 
corrosion potential, natural productivity, water impoundment res- 
ervoir embankments, and trafficability. 

Each criterion is subdivided into a range. For example, topog- 
raphy slope ranges are subdivided into the following percentages: 
to 2, 2 to 5, 5 to 12, 12 to 25, and in excess of 25. The individual 
soils delineated in detailed soil surveys may now be placed into the 
above topography ranges according to their slope gradients. The 
different slopes are assigned a class name depending on the use for 
which the soil is being evaluated. Verbal rating terminology for 
classes has not been standardized but there is a similarity in the 
following terms: 

Very Good Most Favorable No Limitations 

Good Very Favorable Some Limitations 

Fair Favorable Moderate Limitations 

Poor Somewhat Unf avorable.__ Severe Limitations 

Very Poor Unfavorable Very Severe Limitations 

A soil within the same slope range may have different ratings 
when considered for different uses. For residential purposes, some 
of the most favorable slopes would be between 2 to 5 per cent be- 
cause of the aesthetic value associated with rolling topography. 
These slopes would only be considered as "good" for natural rec- 
reation. On the other hand, 12 to 25 per cent slopes are considered 
to have "moderate limitations" for residential use but are very good 
for natural recreation. Slopes in excess of 25 per cent are consid- 
ered "unfavorable" for residences and have "severe limitations" 
when used for transportation. 

Ratings of the criteria are generally made for multipurpose 
use. For urban planning this includes land used for residence, rec- 
reation, industry, and transportation. The residential land use is 
further subdivided into areas using community sewage disposal 



142 Quarterly Journal of the Florida Academy of Sciences 

systems and areas using septic tank disposal systems. Recreation 
is commonly subdivided into the following two categories: devel- 
oped (such as golf courses) and natural (such as picnic or camp 
sites). Predictions concerning the behavior of a soil for any of 
the above specified uses may be made after the soil qualities have 
been determined. 

Permeability is a quality of the soil that enables it to transmit 
water. Soils with moderately slow to very slow permeability are 
not suited for estate type residences that are dependent on septic 
tanks for sewage disposal. The effluent absorption rate is too slow 
for septic tanks to function properly. It is too late for a soil sur- 
vey to help with soil interpretations of permeability where houses 
have already been built and septic tanks are failing! Slow perme- 
ability would not affect the development of residential areas with 
community-type sewage disposal systems. The suitabilities of a 
few widely occurring soils of Florida for septic tank sewage dis- 
posal systems are shown in Table 1. Similar tables, based on soil 

TABLE 1 

Suitability of soils for septic tank sewage disposal systems* 







Principal 


Soil 


Rating 


Limitations 


Lakeland fine sand, 






to 2 per cent slopes 


Very Good 


None 


Lakeland fine sand, 






5 to 12 per cent slopes 


Very Good 


None 


Leon fine sand, 






to 2 per cent slopes 


Fair 


Wetness 


Rutlege fine sand, 






to 2 per cent slopes 


Very Poor 


Wetness 


Weston fine sand, 






to 2 per cent slopes 


Very Poor 


Wetness and 


Norfolk loamy fine sand, 




slow permeability 


to 2 per cent slopes 


Fair 


Permeability 


Norfolk loamy fine sand, 






5 to 12 per cent slopes 


Fair 


Permeability 


Everglades muck 


Very Poor 


Wetness 



Without modification of natural conditions. 



Carlisle: Soils Survey for Planning Urban Development 143 

qualities and characteristics, may be constructed to predict the be- 
havior of soils for other uses. 

Wetness or depth to the ground water table can seriously affect 
the suitability of a soil for most uses. Areas subject to overflow 
or flood are closely associated with wetness. Soil maps show the 
intricate drainage pattern of an area and provide an excellent basis 
for planning all types of water control measures. Construction 
of buildings, roads, railroads, and airfields is very severely limited 
on wet soils as good drainage is required to attain maximum bear- 
ing capacity of the soil. Drainage of wet soils and soils with sea- 
sonably high water tables is essential for septic tank filter fields to 
function properly. Good drainage is also desirable for camp sites 
and picnic areas. On the other hand, wet areas provide better 
refuge for wildlife and may be well-suited for game reserves and 
hunting areas. 

Depth to bedrock is important in most construction work. Soils 
with bedrock less than one foot below the surface are seriously 
limited for many uses. The cost of laying cables, gas and oil pipe- 
lines, and sewer mains may sky-rocket if bedrock is encountered. 
Septic tank installations are expensive and may not function prop- 
erly. Through use of soil surveys, utility pipelines may be planned 
to follow routes that are relatively free of rock. Areas where bed- 
rock occurs close to the surface may be utilized for recreational 
purposes. 

Shrinkage and swelling of soils with changes of moisture content 
depend primarily on the kind and amount of clay present. Soils 
with high contents of expanding clays will swell when wet and 
shrink when dry. The stresses created by volume changes of this 
type may be great enough to break cast-iron utility pipes. Volume 
changes are frequently responsible for cracked walls in private and 
public buildings. Foundations crack or slip and roads may buckle 
on soils with high contents of expanding clays. Since soil maps 
show areas with likely shrink-swell problems, they may be used 
to locate more suitable building sites or to anticipate the need 
of costly overdesign in construction. 

Bearing values, which reflect the ability of a soil to sustain 
static or mobile loads, are used by engineers to determine the 
stability of soils for building foundations. Organic soils are poorly 
suited for construction because of their low ability to support loads. 
Soil materials that are high in organic deposits must be removed 



144 Quarterly Journal of the Florida Academy of Sciences 

and replaced by more stable materials at considerable cost. Soils 
of high organic content are delineated on soil maps. 

Erosion of soil may be a serious problem in urban-fringe areas. 
This soil characteristic is important in the construction of dwellings, 
roads, airfields, and light industries. Sloping areas where the pro- 
tective vegetation has been removed are particularly susceptible. 
As development progresses there is less soil exposed to absorb a 
greater amount of water which is concentrated by roofs, pave- 
ments, and other structures. The sediments from an eroded area 
may be repeatedly washed into streets, filling ditches and obstruct- 
ing culverts. Plans for the control of erosion and deposition of 
sediments by proper disposal of excess water may be easily de- 
veloped through the interpretation of soil maps. 

Corrosion of metals is usually explained by the connection of 
dissimilar metals, junction of dissimilar soils, reaction of soils, bac- 
terial activity, and stray currents. Each dissimilar soil junction sets 
up a current flow with the resulting amount of corrosion depending 
on the electrical resistivity of the soil. Ranges of soil reaction may 
be easily plotted on soil maps. Frequently, the amount of oxida- 
tion caused by sulfate-reducing anaerobic bacteria may be corre- 
lated with certain soil types. Soil maps may be used to pinpoint 
areas of dissimilar soils and to outline soils in which a high degree 
of sulfate-reducing bacterial activity is likely to occur. 

Natural productivity affects the suitability of soils for recrea- 
tion, transportation, and residential uses. Most productive soils 
are ideal for growing a large variety of plants. Productive soils 
offer a greater choice of grass varieties for lawns or golf courses 
than unproductive soils. Road shoulders are stabilized more rap- 
idly and successfully where productive soils occur or suitable 
topsoil is used. Soils with poor structure may be improved by 
addition of organic matter. The amount of wildlife is frequently 
affected by the natural productivity of soils. 

Water impoundment is needed in residential areas using lagoon- 
type sewage disposal and is frequently desirable in constructing 
small ponds. The ponds may be stocked with fish or used only 
for water storage. Small ponds add to the aesthetic value of golf 
courses and are frequently used as a source of water for irrigation. 
Soils vary widely in their suitability for impounding water. 

Trafficability refers to the ease of passage over an area by 
foot or light vehicle. It may determine the suitability of an area 



Carlisle: Soils Survey for Planning Urban Development 145 

to be used for hunting, picnic sites, and camp sites. The better 
drained soils are usually more favorable than the poorer drained 
soils. This soil quality is of significance only in determining the 
limitations of soils for recreational use. 

After a soil is evaluated for all of the above characteristics or 
qualities; it may be assigned an over-all rating as to its suitabilities 
or limitations for residential, industrial, transportational, and rec- 
reational use. Multipurpose suitabilities for a few widely occur- 
ring soils of Florida are shown in Table 2. These suitabilities are 
based on natural soil properties that limit the soils for a particular 
use. This does not mean that soils rated as "poor" or "very poor" 
cannot be developed for that particular use; however, it does mean 
that problems will be encountered in overcoming the severe limi- 
tations of these soils for the use being considered. 

Houses can be built on soils rated as "very poor" for residential 
purposes and roads can be constructed on soils rated as "very 
poor" for transportation; however, the cost of overdesign is very 
high. Frequently the cost is too great for residential construction 
to be economically feasible. In some instances, facilities must be 
constructed on a given location without regard to the soils involved. 
Interpretation of the soil map will show the kind of limitations 
present and the degree of treatment or overdesign needed to over- 
come these limitations. 

For interpretative purposes, soils with similar predicted be- 
haviors may be grouped together. Colored maps are frequently 
used for comprehensive planning as an aid in depicting the various 
soil qualities and characteristics. Colored interpretative maps de- 
lineating the suitability of soils for specific uses enable the plan- 
ning official to present the facts to an inquirer while he is still in- 
terested and before he makes a decision. Although soil survey 
interpretations serve as a useful adjunct to urban planning, it 
should be emphasized that there is no substitute for on-site investi- 
gations. 

Before the detailed soil survey can be used effectively in urban 
planning, a map of the area should be constructed depicting the 
current land use. This map is usually made on acetate overlays 
superimposed on aerial photographs having an identical scale as 
the detailed soil maps. The overlays show such information as 
use of the land for range, forest, crops, idle land, swamps, water, 
roads, railroads, electrical power lines, gas lines, houses, businesses, 



146 



Quarterly Journal of the Florida Academy of Sciences 



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Carlisle: Soils Survey for Planning Urban Development 147 

public buildings, and industrial buildings. The completed map 
presents a concise representation of land use within the area and 
should be revised periodically as urban development progresses. 

Summary 

Detailed soil surveys are serving officials in many areas through- 
out the country as a basis for planning urban development. The 
soils of any area may be evaluated in terms of soil qualities and 
characteristics. By using this information, soils may be rated ac- 
cording to their suitabilities for residential, industrial, transporta- 
tional, and recreational use. After tentative plans for the use of 
an area have been made, there is no substitute for on-site investi- 
gations. Soil surveys are put to their best use in the urban fringe 
when engineers, architects, planners, and soil scientists work co- 
operatively with local county officials in order to design a unified 
plan of urban development. 

Literature Cited 

Bartelli, L. J. 1962. Use of soils information in urban-fringe areas. Jour. 
Soil and Water Cons., vol. 17, no. 3, pp. 99-103. 

Clayton, J. W., H. Kennedy, H. C. Porter, and R. E. Devereaux. 1959. 
Use of soil survey in designing sewage disposal systems. Virginia Agr. 
Exp. Sta. Bull. 509, 15 pp. 

Kellogg, C. E. 1955. Soil surveys in modern farming. Jour. Soil and 
Water Cons., vol. 10, no. 6, pp. 271-277. 

Klingebiel, A. A. 1963. Land classification for use in planning. USDA 
Yearbook of Agriculture, pp. 399-407. 

Marshall, R. M. 1963. Soil surveys are guides for many urban-area de- 
velopments. Soil Cons., vol. 29, no. 4, pp. 75-77. 

Obenshain, S. S., H. C. Porter, and R. E. Devereaux. 1962. Soil survey 
for urban planning and other uses. Virginia Agr. Exp. Sta. Bull. 538, 
26 pp. 

Soil Survey Staff. 1951. Soil Survey Manual. USDA Agr. Handbook No. 
18, 503 pp. 

Department of Soils, University of Florida, Gainesville, Florida. 
Quart. Jour. Florida Acad. Sci. 27(2) 1964 



DEMOGRAPHY OF A FLORIDIAN ALCOHOLIC SAMPLE 
James H. Williams 

The intent of the writer is to describe certain characteristics of 
an alcoholic sample and compare them with the population of the 
state in which they reside. These characteristics include the demo- 
graphic factors of age, sex, marital status, education, veteran status, 
employment status, and level of last occupation. 

The sample includes the 941 patients admitted to the State of 
Florida Alcoholic Rehabilitation Center during a consecutive six- 
teen month period of 1962-63. This Center is located in Avon 
Park and provides inpatient treatment facilities for state residents. 

Data on the Florida population were taken from the 1960 
Census of Population. Included were the comparable 2,002,476 
urban white persons 20 years of age or over enumerated in Flor- 
ida during 1960. 

Age and Sex 

In the alcoholic sample, males constituted the overwhelming 
majority of the subjects in comparison with females. Almost three- 
fourths (72 per cent) of the alcoholics were males and slightly over 
one-fourth (28 per cent) females. The comparable Florida popu- 
lation consisted of only 47 per cent males and 53 per cent females. 
Thus, the alcoholic males had a difference in per cent of 25 over 
the state. 

This difference in per cent and all others mentioned in the 
paper, unless otherwise stated, were significant at or beyond the .05 
level. The statistical test was the significance of difference be- 
tween proportions (Hagood and Price, 1952). 

When comparing sex differences between the alcoholic sample 
and the population of Florida by three age groups, the highest level 
of contrast occurred in the older age group, 55 years of age or 
over. The percentage of the state's male population remained 
fairly constant for the younger ages (20-24), the middle group 
(35-54), and the older males (55 or over), while the percentage of 
males in the alcoholic sample was remarkably higher in the older 
age group. Thus, while the younger and middle age groups of 
alcoholics had from 70 to 72 per cent male, the older age group 
had 86 per cent male. The difference in percentages between the 



Williams: Demography of Alcoholics 



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150 Quarterly Journal of the Florida Academy of Sciences 

sample and state population ranged from 24 among younger ages 
to 22 in the middle to 39 in the older age group. 

Persons in the alcoholic sample, in the main, were between 
ages 35 and 55 years old. The alcoholic males had 71 per cent 
and the females 80 per cent of their cases in this age range com- 
pared with the Florida males at 39 per cent and females at 38 
per cent. The differences in per cent between the alcoholic sam- 
ple and the state population were 32 and 42 for males and females, 
respectively. The mean age for the male alcoholic was 45.82 
years, somewhat younger than the comparable figure for Florida 
males with a mean of 46.87. In the female alcoholic sample the 
mean age was 44.04 compared with 47.31 in the state population. 
The difference in means was 1.05 years for males and 3.27 for 
females. Both differences in means showed the alcoholic sample 
to be younger than the Florida population. 

An illustration of age-sex composition of the sample and popu- 
lation is presented in Fig. 1. This type of graph is based on the 
percentages of all persons in the sample who were in each age- 
sex group. The sum of percentages in the sample or the popula- 
tion equals one hundred for both males and females combined. 
This technique makes possible not only comparisons between the 
sample and the population but also the relative contribution of 
each sex to these two groups. 

Among males the percentage of alcoholics in each age group 
exceeded the state population beginning at ages 30-34, ascending 
at 35-39, reaching a summit at 40-44, and descending in each sub- 
sequent group to fall below the population base for each of the 
last two age categories, 60-64 and 65 or over. On the female side 
of the graph, there were no alcoholic cases that fell below age 25. 
Beginning with age 25 there was a steady increase, with the 40-44 
age group surpassing the base population. The 45-49 age group 
rose just slightly above the Florida population and the 50-54 
dropped slightly below. There was a sharp decline in the remain- 
ing age groups, 55-59, 60-64, and 65 or over. 

The predominance of male over female alcoholics stands out 
boldly when we once again observe Fig. 1. Thus, in the alcoholic 
sample, in no age group was the percentage greater for women 
than for men. However, the shapes of the graph for men and 
women are similar in that the largest proportions for both sexes 
occurred in the 40-44 age group. 



Williams: Demography of Alcoholics 151 

Marital Status and Number Times Married 

The extent of marital stability of alcoholics was of considerable 
interest for this report. The two indices of this stability used were 
marital status and number of times ever married. 

To be considered first is marital status. About 50 per cent 
of the male alcoholics and 42 per cent of the female alcoholics 
reported their marriages intact at treatment intake. The compa- 
rable per cent of Florida males married at time of the census 
enumeration was 82 and for females the per cent was 72. Differ- 
ences in per cent of persons married between the alcoholic sample 
and the state population were 32 among males and 30 for females. 
Thus, even though significantly greater proportions of persons in 
the state population were married than was the case in the alco- 
holic sample it should be noted that the largest percentage of sub- 
jects in both of these groups were in the most stable category, 
married. 

Second in magnitude for the alcoholic sample was the least 
stable categories, divorced or separated. At treatment intake, 27 
per cent of the male alcoholics and 25 per cent of the female alco- 
holics were divorced. Only 3 per cent of the state males and 5 
per cent of the females were divorced at the time of the 1960 Cen- 
sus of population. The differences in per cent of persons divorced 
in the alcoholic sample and in the state were 24 for males and 20 
for females. The male alcoholics had 15 per cent of their num- 
ber who were alienated by separation from their spouses while 
females showed 14 per cent compared with only 1 and 2 per cent 
of the respective state groups. Thus, on both of the above meas- 
ures of alienation in the marital relationship the alcoholics ex- 
ceeded the state standard. 

The two remaining marital status categories were never mar- 
ried and widowed. In the alcoholic sample only 6 per cent of 
the males and 5 per cent of the females had never married. These 
figures not only were relatively low in the alcoholic group but they 
dropped below the state figures. Only among males was the differ- 
ence between the alcoholic sample and the state statistically sig- 
nificant. Those persons widowed constituted only 2 per cent of 
male alcoholics and 14 per cent of female alcoholics. No signifi- 
cant difference was found between the per cent of alcoholics wid- 
owed and the per cent of the Florida population in this category. 



152 Quarterly Journal of the Florida Academy of Sciences 

A similar pattern of relationship between alcoholism and marital 
status remained when the three age groups were observed. Thus, 
in all six age and sex groups, significantly fewer alcoholics were 
married while at the same time more of them were divorced or 
separated than was true in the state population. 

Upon closer observation of the data, the alcoholic males in the 
sample showed an increase from 40 per cent married in the 
younger age group (20-34) to 49 per cent in the middle group 
(35-54) to 59 per cent in the older ages (55 or over). The Florida 
population did not show a corresponding increase with age. The 
females in the alcoholic sample showed a decrease in proportion of 
those married as age increased. Fifty-five per cent of the younger 
females, 41 per cent of the middle age ones, and only 32 per cent 
of the ones who were in the older group were married at treatment 
intake. The comparable percentages of married females in Flor- 
ida were 82 in both the younger and middle groups but dropped 
to 56 per cent married in the older one. 

When the divorced and separated statuses were combined into 
one index of marital instability, interesting profiles appeared. 
Among male alcoholics that were divorced or separated, 45 per 
cent fell in the younger age group, 44 per cent in the middle group, 
and a decline to 30 per cent in the older age group. In the female 
sample the largest proportion was in the middle age group (40 
per cent) with the younger age group at 36 per cent and the older 
at 21 per cent. The state population did not follow the patterns 
set by the alcoholic sample. 

Among alcoholic women the proportion widowed in the middle 
and older age groups was not only relatively high but also was sig- 
nificantly higher than the state population. Thus, 3 per cent of 
alcoholic women in the younger age group were widowed, 14 per 
cent in the middle group, and 42 per cent in the older age group 
were widowed. The state proportions widowed showed a similar 
climb by age but at a lower level. 

Marital stability is further indicated by observing the total num- 
ber of times the subjects have ever been married. This gives us 
a crude index of the history of marital stability or instability in the 
alcoholic sample when compared with its state counterpart. At 
treatment intake, the majority of the alcoholic males (56 per cent) 
and a large proportion (42 per cent) of the females had married 
once. Of the state population reporting one marriage, the males 



Williams: Demography of Alcoholics 153 

had 71 per cent and the females 74 per cent. The difference in 
the two groups was 15 per cent for males and 32 per cent for 
females. 

Male alcoholics appeared to be more stable on this index than 
females. Thirty-nine per cent of males compared with 53 per 
cent of females had been married two or more times. The propor- 
tion of persons with two or more marriages for the state was be- 
tween 19 and 20 per cent for both males and females. Thus, the 
above data would suggest that male alcoholics in this sample 
showed more evidence of greater marital stability than females. 
Such a difference was not indicated in the state population. 

When number of marriages was set out by the six age and sex 
groups, the above mentioned differences remained. The alcoholic 
sample showed fewer persons in each age and sex group who had 
one marriage. More alcoholics reported two or more marriages 
than the comparable Florida population. Finally, male alcoholics 
in every age group showed more marital stability than female alco- 
holics. This pattern was not followed in the Florida population. 

Educational Attainment 

No consistent over-all difference was found in the educational 
level between the alcoholic sample and its Florida counterpart. 
Males in the sample had completed on the average of 10.94 years 
of school while females had 11.62 years. No significant difference 
was found between alcoholic males and those in the state popula- 
tion. Among females the alcoholics were just slightly higher than 
the Florida group. 

Upon observation of the alcoholic sample in relation to educa- 
tion the following was found: about one-fourth of both males and 
females had some college; 28 per cent of males and 36 per cent 
of females had completed high school; 21 per cent of males and 24 
per cent of the females had some high school; and 26 per cent 
of males and only 15 per cent of females had grade school or less. 

Veteran Status 

Census data were not available on females by veteran status. 
However, only 7 per cent of the sample of alcoholic women were 
veterans. On males, the census data were not available for urban 



154 Quarterly Journal of the Florida Academy of Sciences 

white males 20 years of age or over as has been used in our other 
statistics but included only white males 25 years of age or over. 

Male veterans in the alcoholic sample out-numbered non-vet- 
erans by a sizeable majority. Sixty-two per cent of the male alco- 
holics were veterans while only 38 per cent of them were non-vet- 
erans. In Florida only 48 per cent of the comparable male popu- 
lation were veterans and 52 per cent were non- veterans. The differ- 
ence in per cent between the alcoholics and state population was 
14 with the alcoholics having significantly more veterans. 

When observed by three age groups, an interesting pattern 
appeared. As age increased in both the alcoholic and Florida 
population, there was a general decrease in proportion of veterans. 
Seventy-eight per cent of the younger age group (25-34) of males, 
66 per cent of the middle one (35-54), and only 31 per cent of the 
older (55 or over) alcoholics were veterans. Sixty per cent of the 
younger, 55 per cent of the middle, and 30 per cent of the older 
males in the state population were veterans. Differences in per 
cent between alcoholic and state males were 14, 10, and less than 
1 in the younger, middle, and older age groups, respectively. The 
older age group dropped below the level of significance. 

Employment Status 

A substantial proportion of the alcoholic sample reported that 
they had been employed just prior to entering alcoholic treatment. 
Forty-four per cent of the male alcoholics were employed and 55 
per cent were unemployed. Among Florida males, 72 per cent 
were employed, according to the Census, and 28 per cent were 
unemployed. The difference in proportion between the alcoholic 
males and the state population was a striking 28 per cent. The 
percentages of employed females in the alcoholic and the state 
group were much lower than males. Of course many females, 
typically, are not in the labor force. Consequently, only 22 per 
cent of the alcoholic females were employed and 78 per cent not 
employed at intake. Among Florida females 32 per cent were 
working and 68 per cent were not working according to the Census 
figures. The difference in per cent between employed alcoholic 
females and those in Florida was 11. 

When age as well as sex was taken into account, the employ- 
ment status of the male alcoholics may be summarized as follows. 



Williams: Demography of Alcoholics 155 

The percentages of male alcoholics employed were: 46 in the 
younger age group; 48 per cent in the middle age category; and 
only 28 per cent in the older age group. In Florida the males had 
90 per cent employed in both the younger and middle age groups 
but only 38 per cent in the older age category. It is further noted, 
that the difference gap in per cent employed between the alcoholic 
sample and the Florida population tended to decrease as age in- 
creased. These differences in per cent ran from 44 in the younger, 
42 in the middle, to only 10 in the older age group. 

The females followed a similar pattern to males except fewer 
of them were employed in every category. Only 9 per cent of 
the younger female alcoholics, 24 per cent of the middle age group, 
and 16 per cent of the older ones were employed. This compared 
with 38 per cent of the younger Florida females, 44 per cent of 
the middle age group, and 16 per cent of the older ones who were 
employed. The percentage gap between alcoholic females and 
comparable state females closed as aged increased. Among young- 
er ones the difference was 29 per cent, 20 percent in the middle, 
and less than 1 for older females. 

Occupational Level of Last Job 

In the distribution of occupational level of last job before treat- 
ment, 33 per cent of the male alcoholics and 23 per cent of the 
females were in the two highest occupational groupings — profes- 
sional, managerial, official, and proprietory. When taken together 
these two occupational groups were not significantly different from 
the comparable state population of employed persons as given in 
the 1960 Census. Persons not in the labor force were excluded 
from both the alcoholic sample and the state population. How- 
ever, among males the alcoholics had fewer professional, technical, 
or kindred workers than the state while this sample had more man- 
agers, officials, and proprietors than the state. No significant differ- 
ence was found between alcoholic females and those in the state 
in these two occupational groups. Eighteen per cent of alcoholic 
males and 42 per cent of females listed clerical or kindred workers 
or sales workers as the last job they held before treatment. No 
significant difference was noted between the alcoholic and state 
groups. 



156 Quarterly Journal of the Florida Academy of Sciences 

Skilled workers — craftsmen, foremen, operatives, or kindred 
workers had 28 per cent among males and 4 per cent for female 
alcoholics. In both sex groups there were fewer alcoholics who 
were skilled workers than was true in the state population. Thus, 
Florida males had 38, while females had only 12 per cent listed as 
craftsmen, foremen, operatives, or kindred workers. 

The final two occupational categories combined turned in the 
opposite direction when alcoholics were compared with Florida 
population. The alcoholic male sample had 21 per cent while 
the labor force female showed 31 per cent either as service workers 
including private household or unskilled laborers. The Florida 
population listed only 12 per cent of the males and 19 per cent 
of its females in these two occupational levels. Thus, more alco- 
holic males and females were service workers or unskilled laborers 
than was true in the state population. 

Further differences appeared when the three age groups were 
used. There was an increase in proportion of male alcoholics in 
the two highest occupational categories as age increased. Twen- 
ty-one per cent of younger persons, 32 per cent of the middle group, 
and 43 per cent of the older age group of alcoholics reported their 
last occupations to be professional, managerial, official, proprietary, 
or kindred workers. A comparable relationship was not observed 
in the female groups. There was not a consistent increase in oc- 
cupational level with age in either state males or females. 

Again for male alcoholics, when all skilled and unskilled oc- 
cupational levels were combined, there was a systematic decrease 
in proportion in these categories as age increased. The younger 
age group had 59 per cent skilled or unskilled workers, the middle 
one 50 per cent, and the older one only 38 per cent. Neither the 
Florida males and females nor the female alcoholics followed 
this trend. 

A limitation on the above discussion of occupational level of 
alcoholics' last job and the occupational level of employed persons 
in Florida according to the 1960 Census of Population should be 
noted. Last job of the alcoholic is not strictly comparable to the 
Census figures. However, they have been included for compari- 
son purposes with definite limitation recognized by the writer. 



Williams: Demography of Alcoholics 157 

Summary 

In this study certain demographic characteristics of an alcoholic 
sample were compared with those of a state population. These 
characteristics included age, sex, marital status, number times ever 
married, education, veteran status, employment status, and level 
of last occupation. 

The sample included 941 patients admitted to the State of 
Florida Alcoholic Rehabilitation Center during a consecutive six- 
teen month period of 1962-63. The state population against which 
the alcoholic sample was compared came from the 1960 Census 
of Population for Florida. Only urban white persons 20 years of 
age or over were included in order to maintain comparability with 
the alcoholic sample. 

Almost three out of four alcoholics in the sample were males 
while less than half of the Florida population were males. The 
overwhelming majority of both male and female alcoholics was in 
the middle age group (35-54), with only minorities of the state 
population falling between these ages. The alcoholic sample in 
both male and female groups was somewhat younger on the aver- 
age than the comparable state population. 

Substantial proportions of the males and females in the alco- 
holic sample were married at treatment intake, but the comparable 
state population had significantly more. The alcoholic sample 
had a higher percentage that were divorced or separated than the 
state. An additional relationship was noted when marital status 
was related to both age and sex. As age increased the alcoholic 
males were more likely to be married. On the other hand, the 
female alcoholics were less likely to be married as age increased. 
The state population did not follow this pattern. 

When total number of marriages was used, a smaller propor- 
tion of both groups of alcoholic males and females had only one 
marriage than did the state population. In turn the alcoholics 
had more persons who reported two or more marriages than the 
comparable state figures. This pattern remained in all three age 
groups. However, male alcoholics in all of these age groups 
showed a greater proportion with two or more marriages than 
female alcoholics. This was not the case in the Florida popula- 
tion. 



158 Quarterly Journal of the Florida Academy of Sciences 

No significant differences were found between alcoholic males 
and Florida males on educational attainment. The average female 
alcoholic had slightly more education than her Florida counter- 
part. 

Male veterans in the sample out-numbered non-veterans by 
a sizeable majority. When the three age groups were used there 
was a systematic decrease in proportion of both alcoholic and 
Florida males who were veterans as age increased. Both the 
younger and middle age groups showed the alcoholics to have a 
greater proportion of veterans than the state. No information was 
available for state female veterans. 

A substantial proportion of the male alcoholics were employed 
just before entering treatment; however, the Florida males re- 
ported a higher proportion. The Florida females showed a higher 
proportion employed than alcoholic females. Females showed 
lower employment percentages than males in both the alcoholic 
sample and the Florida population. 

In the two highest occupational groupings — professional, mana- 
gerial, official, and proprietory — when taken together there was no 
significant difference between the alcoholic sample and the state 
population. Nor was there a difference between these groups of 
persons who were clerical, sales, or kindred workers. However, 
in both sex groups there were fewer alcoholics who were craftsmen, 
foremen, operatives, or kindred workers than in the state popula- 
tion. On the other hand there were more alcoholics than state 
persons who were service or unskilled laborers on their last jobs. 

Acknowledgment 

Data for this paper were obtained in connection with a three- 
year research and demonstration project entitled: "Analysis and 
Evaluation of Collaborative Treatment of Selected Alcoholics." 
This larger investigation was supported, in part, by a research 
grant, Number RD-640, from the Vocational Rehabilitation Admin- 
istration, Department of Health, Education, and Welfare, Wash- 
ington, D. C. The larger project was sponsored by the State of 
Florida Alcoholic Rehabilitation Program, Avon Park, Florida, in 
cooperation with the Division of Vocational Rehabilitation, State 
Department of Education, Tallahassee, Florida. 



Williams: Demography of Alcoholics 159 

Literature Cited 

Hagood, M. J., and D. O. Price. 1952. Statistics for Sociologists. Henry 
Holt and Company, New York, pp. 315-320. 

U. S. Department of Commerce, Bureau of the Census. United States 
Census of Population: 1960, Florida. 

U. S. Government Printing Office. Detailed Characteristics, Final Report 
PC(1)-11C, Table 37, p. 11-134. PC(1)-11D, Table 103, pp. 11-329— 
11-331. Table 104, p. 11-339. Table 105, pp. 11-341, 11-342, 11-344. 
Table 115, pp. 11-396—11-398. Table 123, pp. 11-487—11-490. 

Project Director, State of Florida Alcoholic Rehabilitation Pro- 
gram, P. O. Box 1147, Avon Park, Florida. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



THE SHRIMP TRACHYPENEUS SIMILIS IN TAMPA BAY 
Carl H. Saloman 

During a faunal sampling project in Florida estuarine waters, 
several species of shrimp were captured along with other crustaceans 
and fishes. Shrimp were particularly numerous at two sampling 
stations during the period August 1961 through December 1963. 
Both stations were in the Tampa Bay system: one in Old Tampa 
Bay and the other at Egmont Key located at the mouth of the Bay. 
Each of these two stations was occupied for 24-hour periods alter- 
nately in consecutive months from August 1962 through December 
1963. Forty-nine other stations in Tampa Bay and one station 
offshore were sampled from August 1961 through December 1963 
during daylight hours only. Types of gear used in the daylight 
studies were trawls, beach seines, and push nets. Trawls were the 
only gear used during the 24-hour studies. 

Numerous specimens of penaeid shrimp were caught during 
the 24-hour biological studies conducted on the east side of Eg- 
mont Key at the mouth of Tampa Bay. The dominant penaeid 
shrimp present was Panaeus duorarum (Burkenroad), along with 
some Trachypeneus constrictus (Stimpson) and Sicyonia typica 
(Boeck). Also present were 97 specimens of Trachypeneus similis 
(Smith). These are the first specimens reported from Tampa Bay 
and the west coast of Florida between Apalachicola Bay and the 
Tortugas grounds. All 97 specimens were caught during darkness 
with either a 10- or 16-foot shrimp try net. No T. similis were 
caught in our regular daylight sampling of Tampa Bay or in 24- 
hour sampling in Old Tampa Bay. Data recorded relative to the 
occurrence of T. similis were size, sex, temperature, and salinity 
(table 1). 

The station at Egmont Key is characterized by a sandy shell 
bottom, a depth of approximately two fathoms of water, and an 
abundant growth of Thallasia testudinum (Konig) and Syringodium 
filiforme (Kutz). 

Discussion 

The month of December was the period of maximum abundance 
of T. similis at the Egmont Key station with 95 of 97 total speci- 
mens being caught during that time. Eldred (1959a) found that 



Saloman: Penaeid Shrimp in Tampa Bay 161 

the period of maximum abundance of T. similis on the Tortugas 
grounds was from January through March 1959 and stated that 
T. similis is next in abundance to P. duorarum on the Tortugas 
grounds, where both species are harvested in the commercial fish- 
ery. Similarly, Eldred (1959b) and Ingle, Eldred, Jones, and Hut- 
ton (1959) found that the major abundance of T. similis occurred 
on the Tortugas grounds from January through April 1958. 

TABLE 1 
Data for Trachypeneus similis from Egmont Key, Florida 





Time 


Number 


Carapace 


Bottom 


Bottom 




of 


and 


Length 


Temperature 


Salinity 


Date 


Capture 


Sex 


mm. 


°C 


o/oo 


8-31-62 


0300 


1 2 


8.7 


30.30 


32.97 


10-30-62 


0300 


1 2 


9.4 


21.32 


31.58 


12-20-62 


2100 


2 $ 


9,3 (8.9-9.6) 


14.40 


32.99 


12-12-63 


2000 


5 $ 
10 2 


7.2 (6.4-7.5) 
9.7 (6.6-12.2) 


16.12 


33.71 


12-12-63 


2300 


13 $ 

32 2 


8.6 (7.5-9.6) 
10.2 (7.5-12.4) 


16.14 


33.80 


12-13-63 


0200 


4 $ 
17 2 


8.0 (6.0-9,3) 
9.8 (6,3-12.2) 


16.37 


33.75 


12-13-63 


0500 


6 $ 
6 2 


8.3 (7,3-8.9) 
8.8 (7.2-10.3) 


16.21 


33.44 


Total 




30 $ 
67 2 


8.26 (6.0-9.6) 
9.85 (6.3-12.4) 







Females averaged larger than males in each of our collections 
containing both sexes. Ingle, Eldred, Jones, and Hutton (1959) 
also found that the females were larger than the males in all the 
monthly summaries of their catch records on the Tortugas grounds. 

All specimens from Egmont Key were found in salinities ex- 
ceeding 31 parts per thousand. 

Burkenroad (1934) reported the distribution of T. similis from 
the Gulf of Mexico, northern Antilles, Venezuela, Louisiana, and 
the western coast of Florida. He stated, however, that T. constric- 
tus inhabits areas north and east of that of T. similis. He also 
stated that the latter replaces T. constrictus on the western and 
southern shores of the Gulf of Mexico and the Caribbean. 



162 Quarterly Journal of the Florida Academy of Sciences 

The occurrence of T. similis throughout the Gulf of Mexico is 
mentioned by various authors. Eldred (1959a, 1959b) and Ingle, 
Eldred, Jones, and Hutton (1959) cited the occurrence of T. similis 
on the Tortugas grounds. Perez-Farfante (1953) mentioned that 
it occurred in the Bahia de Cienfuegos of Cuba. Hildebrand (1954) 
in his study of the fauna of the brown shrimp (Penaeus aztecus Ives) 
grounds in the western Gulf of Mexico found that it was second to 
P. aztecus in abundance. He also noted that T. similis was uncom- 
mon in depths of less than 12 fathoms and appeared to prefer a 
mud bottom. Guest (1956) mentioned that T. similis occurs in bays 
and the Gulf of Mexico along the Texas coast. Renfro and Brusher 
(1963) in their work on shrimp populations between Galveston, 
Texas, and Cameron, Louisiana, found the majority of T. similis 
occurring in 7Vfe and 15 fathoms, but some in depths as great as 45 
fathoms. Burkenroad (1939) listed the species as occurring in 
Pensacola Bay, Florida, and stated that it is a frequently encount- 
ered species west of 88° 10' in the Gulf of Mexico. He also found 
that T. similis appeared to prefer a muddy bottom. Wass (1955) 
did not mention its occurrence in his samples from Alligator Harbor 
and adjacent inshore areas of northwestern Florida but stated that 
it is known to occur off the coast of northwestern Florida within the 
30-fathom line. 

The omission of this species from several papers on the fauna 
of the Gulf waters of Florida is notable. In a bait shrimp survey 
of the west coast of Florida Woodburn, Eldred, Clark, Hutton, 
and Engle (1957) did not mention the occurrence of T. similis. 
When reporting on the pink shrimp, Penaeus duorarum, in Florida 
waters, Eldred, Ingle, Woodburn, Hutton, and Jones (1961) did not 
list this species. T. similis was also absent from the lists of Hutton, 
Eldred, Woodburn, and Ingle (1956), reporting on a study of the 
ecology of Boca Ceiga Bay, Florida. Eldred (personal communica- 
tion) found this species only on the Tortugas grounds and in Pen- 
sacola and Apalachicola Bays on the west coast of Florida. Pub- 
lished research results by Tabb and Manning (1961) from northern 
Florida Bay and by Tabb, Dubrow, and Jones (1962) from Ever- 
glades National Park, Florida, did not mention the occurrence of 
T. similis. Nevertheless, the record of T. similis from Tampa Bay 
now shows that its distribution probably extends throughout the 
entire periphery of the Gulf of Mexico. 



Saloman: Penaeid Shrimp in Tampa Bay 163 

Literature Cited 

Burkenroad, M. D. 1934. The Penaeidae of Louisiana with a discussion on 
their world relationships. Bull. Amer. Mus. Nat. Hist., vol. 68, art. 2, 
pp. 61-143. 

. 1939. Further observations on Penaeidae of the northern Gulf of 

Mexico. Bull. Bingham Oceanogr. Coll., vol. 6, no. 6, pp. 1-62. 

Eldred, Bonnie. 1959a. Notes on Trachypeneus (Trachysalambria) similis 
(Smith) in the Tortugas shrimp fishery. Quart. Jour. Florida Acad. 
Sci., vol. 22, no. 1, pp. 75-76. 

. 1959b. A report of the shrimps (Penaeidae) collected from the Tor- 
tugas controlled area. Florida State Board Conserv., Spec. Sci. Rept., 
no. 2, pp. 1-6. 

Eldred, B., Robert M. Ingle, Kenneth D. Woodburn, Robert F. Hutton, 
and Hazel Jones. 1961. Biological observations on the commercial 
shrimp, Penaeus duorarum Burkenroad, in Florida waters. Florida State 
Board Conserv., Prof. Pap. Ser., no. 3, pp. 1-139. 

Guest, W. C. 1956. The Texas shrimp fishery. Texas Game and Fish 
Comm. Bull., no. 36, pp. 1-23. 

Hildebrand, H. H. 1954. A study of the fauna of the brown shrimp (Pen- 
aeus aztecus Ives) grounds in the western Gulf of Mexico. Publ. Inst. 
Mar. Sci. Univ. Texas, vol. 3, no. 2, pp. 233-366. 

Hutton, R. F., B. Eldred, K. D. Woodburn, and R. M. Ingle. 1956. The 
ecology of Boca Ceiga Bay with special reference to dredging and fill- 
ing operations. Florida State Board Conserv., Mar. Lab., Tech. Ser., 
no. 17, pp. 1-87. 

Ingle, R. M., B. Eldred, H. Jones, and R. F. Hutton. 1959. Preliminary 
analysis of Tortugas shrimp sampling data, 1957-58. Florida State 
Board Conserv., Tech. Ser., no. 32, pp. 1-45. 

Perez-Farfante, Isabel. 1953. The commercial shrimp of Cuba. Memoirs 
of the Cuban Society of Natural History, vol. 21, no. 2, pp. 1-44. 
(Translated by Petronila C. Prado, Gulf Fishery Investigations, Fish 
and Wildlife Service). 

Renfro, William C, and Harold A. Brusher. 1963. U. S. Fish and 
Wildlife Service. Bur. Comm. Fish., Cir. 161, pp. 13-17. 

Tabb, Durbin C, David L. Dubrow, and Andrew E. Jones. 1962. Studies 
on the biology of the pink shrimp, Penaeus duorarum Burkenroad, in 
Everglades National Park, Florida. Florida State Board Conserv., Tech. 
Ser., no. 37, pp. 1-32. 



164 Quarterly Journal of the Florida Academy of Sciences 

Tabb, Durbin C, David L. Dubrow, and Raymond B. Manning. 1962. 
The ecology of Northern Florida Bay and adjacent estuaries. Florida 
State Board Conserv., Tech. Ser., no. 39, pp. 1-81. 

Tabb, D. C, and R. B. Manning. 1961. A checklist of the flora and fauna 
of northern Florida Bay and adjacent brackish waters of the Florida 
mainland collected during the period July, 1957 through September 
1960. Bull. Mar. Sci. Gulf and Carib., vol. 11, no. 4, pp. 552-649. 

Wass, M. L. 1955. The decapod crustaceans of Alligator Harbor and ad- 
jacent inshore areas of northwestern Florida. Quart. Jour. Florida 
Acad. Sci., vol. 18, no. 3, pp. 129-176. 

Woodburn, Kenneth D., Bonnie Eldred, Eugene Clark, Robert F. Hut- 
ton, and Robert M. Ingle. 1957. The live bait shrimp industry of 
the west coast of Florida (Cedar Key to Naples). Florida State Board 
Conserv. Mar. Lab., Tech. Ser., no. 21, pp. 1-33. 

Bureau of Commercial Fisheries Biological Station, St. Peters- 
burg, Beach, Florida. Contribution No. 11. 



Quart. Jour. Florida Acad. Sci. 27(2) 1964 



FLORIDA ACADEMY OF SCIENCES 



COUNCIL FOR 1964 

President: George K. Reid 

President Elect: O. E. Frye, Jr. 

Secretary: John D. Kilby 

Treasurer: John S. Ross 

Past President: Alfred P. Mills 

Past President: Alex. G. Smith 

Chairman, Charter and By-Laws Committee: Elmer C. Prichard 
Finance Committee: James B. Fleek 
Honors Committee: Clarence C. Clark 
Quarterly Journal Committee: I. C. Foster 
State Coordinating Committee: O. E. Frye, Jr. 
Talent Search Committee: Earl D. Smith 
Program Committee: John D. Kilby 
Resolutions Committee: I. G. Foster 

Editor, Quarterly Journal: Pierce Brodkorb 

Chairman, Biological Sciences Section: Casimer T. Grabowski 
Physical Sciences Section: Jackson P. Sickels 
Social Sciences Section: Bernard Bonniwell 
Medical Sciences Section: Veronica Armaghan 
Science Teaching Section: F. S. Shuttleworth 
Conservation Section: Howard R. Bissland 

AAAS Council and Conference Representative: John D. McCrone 

State Coordinator of the Junior Academy: Louise V. Ash 

Councilor at Large, Elected: Elmer C. Prichard 

Councilor at Large, Elected: Glen E. Woolfenden 

Councilor at Large, Appointed: Ruth S. Breen 

Councilor at Large, Appointed: J. E. Hutchman 



ADDITIONAL COMMITTEE CHAIRMEN 

Auditing: James B. Fleek 
Awards and Grants: Alfred H. Lawton 
Future Annual Meetings: Margaret Gilbert 
Local Arrangements: Ruth S. Breen 
Membership: O. E. Frye, Jr. 



166 Quarterly Journal of the Florida Academy of Sciences 

Necrology: D. C. Swanson 

Nominating: Alfred P. Mills 

Visiting Scientist Program: Paul A. Vestal 



MEMBERS OF THE ACADEMY 

May 31, 1964 

Sectional membership is indicated as follows: B, Biological 
Sciences; C, Conservation; M, Medical Sciences; P, Physical Sci- 
ences; S, Social Sciences. Members are requested to inform the 
Secretary of their sectional preferences. 

Agens, Frederick F., 406 Mission Hills Avenue, Tampa 10, Florida P 

Akin, Benjamin, 1019 N. W. 11th Court, Miami 36, Florida B 

Albertson, Mary Susan, 4354 Gun Club Road, West Palm Beach, Florida B 

Alexander, Dr. Taylor R., Botany Dept., Univ. Miami, Coral Gables, Florida B 

Allabough, Edwin, Jr., 250 N. W. Cerritos. Palm Springs, California B 

Allen, D. E., Univ. South Florida, Tampa, Florida 

Allen, Francis R., 2236 Ellicott Drive, Tallahassee, Florida S 

Allen, Dr. John S., President, Univ. South Florida, Tampa, Florida P 

Almodovar, Dr. Luis R., Box 286, San German, Puerto Rico B 

Alt, Dr. David, Dept. Geology, Univ. Florida, Gainesville, Florida P 

Anderson, Dr. William D., Jr., Bureau of Commercial Fisheries, Biological 

Lab., P. O. Box 280, Brunswick, Georgia B 
Andrews, Donald H., 750 N. E. 33rd Street, Boca Raton, Florida 
Andrews, Mrs. Dorothy C, Drawer 959, Brewster Hall, Bradenton, Florida S 
Arean, Dr. Victor M., Dept. Pathology, Univ. Florida, Gainesville, Florida M 
Armaghan, Dr. Veronica, 6430 S. W. 42nd Terrace, Miami 43, Florida B 
Arnold, Dr. Luther A., 143 Norman Hall, Univ. Florida, Gainesville, Florida B 
Ash, Mrs. Louise V., 2405 N. W. 18th Place, Gainesville, Florida T 
Ashford, Dr. Theodore A., Univ. South Florida, Tampa, Florida B 
Auffenberg, Walter, Florida State Museum, Gainesville, Florida B 
Ballard, Dr. Stanley S., Dept. Physics, Univ. Florida, Gainesville, Florida P 
Banks, Joseph E., 6811 Capilla Street, Coral Gables, Florida P 
Barkuloo, James M., P. O. Box 366, Crescent City, Florida 
Barrett, Mary F., 70-B Fremont St., Bloomfield, New Jersey B 
Barton, Dr. Maurice A., 1900 Almeria Way S., St. Petersburg, Florida M 
Batty, David H., 9541 Holiday Road, Miami 57, Florida 
Beck, William M., Jr., 1621 River Bluff Rd., Jacksonville 11, Florida B 
Becknell, Dr. G. G., 6900 Dixon Avenue, Tampa 4, Florida P 
Beery, Dr. John R., Dean, School of Education, Univ. Miami, Coral Gables, 

Florida S 
Bellamy, Dr. R. E., 2311 B Street, Bakersfield, California B 
Bellomy, Mildred D., 80 E. 53rd Terrace, Hialeah, Florida 



Membership List 167 

Berry, Frederick H., U. S. Fish and Wildlife Service, P. O. Box 280, Bruns- 
wick, Georgia B 
Bieber, Theodore I., Dept. Chemistry, Florida Atlantic Univ., Boca Raton, 

Florida P 
Bird, L. C, 303 S. 6th Street, Richmond, Virginia B 

Birdsey, M. R., Dept. Biology, Miami-Dade Junior College, Miami, Florida B 
Birkenholz, Dr. Dale E., Dept. Biological Sciences, Illinois State University, 

Normal, Illinois B 
Bissland, Howard R., 2162 N. W. 23rd Blvd., Gainesville, Florida B 
Bianchard, Dr. Frank N., Dept. Geology, Univ. Florida, Gainesville, Florida P 
Block, W. F., 120 S. W. 25th Street, Gainesville, Florida P 
Bonninghausen, Russel A., 313 E. Lakeshore Dr., Tallahassee, Florida 
Bonniwell, Bernard, Univ. Villanova, Villanova, Penn. 
Boss, Dr. Manley L., Florida Atlantic Univ., Boca Raton, Florida B 
Boulware, Joe W., Univ. South Florida, Tampa, Florida 
Boyd, Dr. Mark F„ 615 East 6th Ave., Tallahassee, Florida B 
Bravos, Mrs. L., Florida State Board of Conservation, Marine Lab., P. O. 

Drawer F, St. Petersburg, Florida 
Breder, Dr. C. M., Jr., American Museum of Natural History, Central Park 

West at 79th Street, New York 24, New York B 
Breen, Dr. Ruth S., Dept. Botany, Florida State Univ., Tallahassee, Florida B 
Brey, Dr. Wallace S., Jr., 1114 N. W. 13th Ave., Gainesville, Florida P 
Brodkorb, Dr. Pierce, Dept. Biology, Univ. Florida, Gainesville, Florida B 
Browder, James Steve, 1716 N. W. 3rd Ave., Gainesville, Florida 
Brown, Dr. Relis B., Dept. Biological Sciences, Florida State University, 

Tallahassee, Florida B 
Broyles, Dr. Arthur A., Dept. Physics, Univ. Florida, Gainesville, Florida P 
Bucher, Charles, 1240 N. W. 203rd St., N. Miami Beach 62, Florida 
Buckland, Charlotte B., 2623 Herschel St., Jacksonville, Florida B 
Bullen, Mrs. Adelaide K., 103 Seagle Bldg., Gainesville, Florida B 
Bullen, Ripley P., 103 Seagle Bldg., Gainesville, Florida B 
Burgess, William D., Florida Christian College, Tampa, Florida B 
Burkman, Ernest, Jr., School of Education, Florida State Univ., Tallahassee, 

Florida S 
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Burns, Russell M., Box 900, Marianna, Florida 
Burton, R. H., 1301 N. W. 180th Terrace, Miami, Florida 
Caldwell, Dr. David K., Los Angeles County Museum, Exposition Park, Los 

Angeles 7, California B 
Campbell, Doak S., 1001 High Street, Tallahassee, Florida S 
Campbell, James A., Box 101, Chipley, Florida B 

Candlin, A. H. S., Div. Science, Jacksonville Univ., Jacksonville, Florida 
Carlisle, Dr. Victor W., 223 McCarty Hall, Univ. Florida, Gainesville, Florida 
Carr, Dr. A. F., Jr., 14 Flint Hall, Univ. Florida, Gainesville, Florida B 
Carr, Joseph A., Jr., 3402 Riverview Drive, Tampa, Florida P 
Carr, Mrs. Marjorie Harris, Micanopy, Florida B 
Carr, Dr. T. D., Dept. Physics and Astronomy, Univ. Florida, Gainesville, 

Florida P 



168 Quarterly Journal of the Florida Academy of Sciences 

Carroll, Dr. Don W., Box 50, Rollins College, Winter Park, Florida P 
Carter, Claude F., 5970 S. W. 46th St., Miami 43, Florida B 
Carver, Dr. W. A., Gold Kist Peanut Growers, Graceville, Florida B 
Chamelin, Dr. I. M., 1923 Elk Ave., Sarasota, Florida B 
Chester, Richard H., Marine Lab., 1 Rickenbacker Causeway, Miami, Flor- 
ida B 
Christensen, Robert Frank, 208 W. 5th Ave., Tallahassee, Florida 
Clark, Dr. Clarence C, Univ. South Florida, Tampa 4, Florida P 
Clark, Samuel F., Dept. Chemistry, Florida Atlantic Univ., Boca Raton, Flor- 
ida P 
Clinton, James H., 206T, Flavet III, Gainesville, Florida 
Cole, Dr. Charles F., Univ. South Florida, Tampa, Florida 
Cole, Mrs. Margaret B., Box 215, Micanopy, Florida 
Conard, Dr. Henry S., Lake Hamilton, Florida P 
Conn, Dr. John F., Box 655, DeLand, Florida P 
Cook, Dr. Thomas, 5515 Ordeena Drive, Coral Gables, Florida B 
Cooke, Samuel H., Roosevelt Junior College, 1235 15th St., West Palm Beach, 

Florida B 
Cooper, Dr. D. M., Ch. 310, Univ. South Florida, Tampa, Florida P 
Coy, Earl H., 60 Sunnyside Dr., Battle Creek, Michigan 
Craig, Dr. Palmer H., Dean of Science and Mathematics, Florida Atlantic 

University, P. O. Box 430, Boca Raton, Florida P 
Craighead, Dr. Frank C, Box 825, Homestead, Florida B 
Creager, Don B., Prof. Biology, Jacksonville Univ., Jacksonville 11, Florida B 
Crist, Dr. Raymond E., Dept. Geography, Univ. Florida, Ganiesville, Florida S 
Cross, Dr. Clark I., Dept. Geography, Univ. Florida, Gainesville, Florida P 
Cunha, Dr. T. J., 252 McCarty Hall, Univ. Florida, Gainesville, Florida B 
Dambaugh, Dr. Luella N., Univ. Miami, Coral Gables 46, Florida S 
Dana, Allan H., R. R. 1, Box 245, Sebring, Florida 
Darlow, Ernest W., P. O. Box 3374, Chamblee, Georgia S 
Davis, Gordon, Motel Farina, 10600 N. W. 27th Ave., Miami, Florida 
Davis, Robert H., Dept. Physics, Florida State Univ., Tallahassee, Florida 
Day, Frederick J., 5515 Maggiore St., Coral Gables 46, Florida 
Day, Orvis E., P. O. Box 74, Greenville, Florida B 
Deichmann, Dr. Wm. B., School of Medicine, Univ. Miami, Coral Gables 34, 

Florida M 
Denman, Dr. Sidney B., Box 1364, Stetson Univ., DeLand, Florida S 
Dequine, John F., Southern Fish Culturists, Box 251, Leesburg, Florida B 
Derr, Dr. Vernon E., 219 N. Lakeland St., Orlando, Florida P 
Devany, Thomas, Marine Lab., 1 Rickenbacker Causeway, Miami, Florida B 
Dew, Dr. Robert J., Jr., Univ. Tampa, Tampa, Florida P 
Dickinson, Dr. J. C, Jr., Florida State Museum, Gainesville, Florida B 
Dickson, John D., Tela Railroad Co., Research Dept., La Lima, Honduras, 

C. A. B 
Dijkman, Dr. Marinus J., 6767 S. W. 112th St., Miami 56, Florida B 
Dobson, Gerard R., Dept. Chemistry, Univ. Georgia, Athens, Georgia P 
Doyle, Dr. Laura M., 365 Hawthorne Avenue, Palo Alto, California 



Membership List 169 

Drach, Mildred A., 360 S. Burnside Ave., Apt. 44, Los Angeles 36, Califor- 
nia S 
Driver, Paul J., 1347 West 9th St., Jacksonville 9, Florida 
Dudley, Prof. Frank M., Div. Physical Sciences, Univ. South Florida, Tampa, 

Florida P 
Dunning, Dr. Wilhelmina F., Dept. Microbiology, Univ. Miami, Coral Gables 

46, Florida B 
Dyrenforth, Dr. L. Y., 3885 St. Johns Ave., Jacksonville, Florida M 
Edwards, Dr. Joshua L., Dept. Pathology, Univ. Florida, Gainesville, Flor- 
ida M 
Edwards, Dr. Richard A., Dept. Geology, Univ. Florida, Gainesville, Flor- 
ida P 
Eickenberg, Charles F., 1347 Sunset Avenue, Lakeland, Florida B 
Elliott, John E., 108 W. 15th St., Apt. 501, Austin, Texas S 
Elson, Herman, 430 Circle Drive, Lake City, Florida 
Emme, Earle E., 1724 Crystal Lake Drive, Lakeland, Florida S 
Eppert, Herbert C, Jr., Dept. Geology, Univ. Florida, Gainesville, Florida P 
Evans, Dr. Elwyn, 601 Magnolia Avenue, Orlando, Florida M 
Ewing, Upton C, 362 Minorca Avenue, Coral Gables, Florida B 
Ferguson, Dr. John C, Florida Presbyterian College, St. Petersburg, Florida 
Fickett, Stephen B., Jr., 404 Highland Avenue, Brooksville, Florida 
Field, Dr. Henry, 3551 Main Highway, Coconut Grove 33, Florida S 
Fitzwater, Dr. Robert N., Dept. Chemistry, Rollins College, Winter Park, 

Florida P 
Fleek, Dr. James B., 518 Patricia Lane, Jacksonville Beach, Florida P 
Fly, Prof. Lillian, 4060 Battersea Road, Coconut Grove, Florida B 
Foote, Dr. Perry A., Coll. Pharmacy, Univ. Florida, Gainesville, Florida P 
Foraker, Dr. Alvan G., 800 Miami Road, Jacksonville 7, Florida M 
Ford, Dr. Ernest S., Dept. Botany, Univ. Florida, Gainesville, Florida B 
Forman, Guy, Dept. Physics, Univ. South Florida, Tampa, Florida P 
Foster, Dr. I. G., Florida Presbyterian College, St. Petersburg, Florida P 
Foster, Dr. Virginia, Box 87, Pensacola College, Pensacola, Florida 
Fox, Dr. Laurette E., P 504, J. Hillis Miller Health Center, Univ. Florida, 

Gainesville, Florida M 
French, Sidney J., Univ. South Florida, Tampa, Florida P 
Friedl, Berthold C, 4931 Riviera Drive, Coral Gables, Florida S 
Friedl, Dr. Frank E., Univ. South Florida, Tampa 4, Florida B 
Froemke, Prof. Robert L., School of Business, Florida State Univ., Tallahas- 
see, Florida S 
Frye, Dr. O. E., Jr., Game and Fresh Water Fish Comm., Tallahassee, Flor- 
ida B 
Fuentes, Joseph, Univ. South Florida, Tampa, Florida 
Fuller, Dorothy L., P. O. Box 418, DeLand, Florida B 

Funderburg, Dr. John B., Jr., Dept. Biology, Florida Southern College, Lake- 
land, Florida B 
Funderburk, E. E., Jr., Box 809, Lake Worth, Florida 
Gallagher, James J., 1005 Keats Ave., Orlando, Florida 



170 Quarterly Journal of the Florida Academy of Sciences 

Garrett, James R., RCA Service Co., Missile Test Proj., Bldg. 989, Mail Unit 

811, Patrick AFB, Florida 
Gathman, C. A., 501 21st Avenue N., Lake Worth, Florida B 
Gilbert, Dr. Margaret, Dept. Biology, Florida Southern College, Lakeland, 

Florida B 
Gilcreas, F. W., Dept. Civil Engineering, Univ. Florida, Gainesville, Florida C 
Gilman, L. C, Dept. Zoology, Univ. Miami, Coral Gables 46, Florida B 
Girard, Murray, 5900 Devanshire Blvd., Coral Gables, Florida B 
Godfrey, Dr. R. K., Dept. Botany, Florida State Univ., Tallahassee, Florida B 
Goethe, C. M., 3731 Tea Street, Sacramento 16, California B 
Goin, Dr. Coleman J., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Golightly, Jacob F., Jacksonville Univ., Jacksonville, Florida 
Grabrowski, Dr. Casimer T., Dept. Zoology, Univ. Miami, Coral Gables, Florida 
Grace, H. T., 12025 Grilling Blvd., Miami 38, Florida S 
Graff, Miss Mary B., Mandarin, Florida S 

Gramling, Dr. L. G., Coll. Pharmacy, Univ. Florida, Gainesville, Florida M 
Gresham, W. B., Jr., P. O. Box 10019, Tampa 9, Florida 
Griffith, Dr. Mildred, Dept. Botany, Univ. Florida, Gainesville, Florida B 
Guilbert, Edward H., Pensacola Junior College, Pensacola, Florida 
Gut, H. J., P. O. Box 700, Sanford, Florida B 
Haas, Mrs. Flora Anderson, R. F. D. 2, Apopka, Florida B 
Haigh, Paul J., Florida Presbyterian College, St. Petersburg, Florida 
Haines, Dr. Charles, P. O. Box Drawer A, Belle Glade, Florida 
Hammett, J. W., P. O. Box 162, Gainesville, Florida 
Hanley, James R., Jr., 6446 Anvers Blvd., Jacksonville 10, Florida 
Hansen, Dr. Keith L., Biology Dept., Stetson Univ., DeLand, Florida B 
Harlow, Richard F., 905 Camphor Lane, DeLand, Florida B 
Harms, Dr. R. H., Poultry Sci. Farm Unit, Univ. Florida, Gainesville, Flor- 
ida B 
Harris, Herbert, Edward Waters College, Jacksonville, Florida P 
Harper, Dr. Roland M., Geological Survey, Univ. Alabama, University, Ala- 
bama B 
Harrington, Dr. Robert W., Jr., P. O. Box 308, Vero Beach, Florida B 
Hatala, Dr. Robert J., 2167 Vivian Way South, St. Petersburg, Florida 
Hazlett, William I., 18610 N. W. 8th Court, Miami 69, Florida 
Hellwege, Dr. Herbert, Rollins College, Winter Park, Florida P 
Hentges, Dr. James F., Jr., 253 McCarty Hall, Univ. Florida, Gainesville, 

Florida B 
Hobbs, Dr. Horton H., Jr., U. S. National Museum, Washington, D. C. B 
Holman, Mrs. B. J., 3320 N. W. 14th Street, Gainesville, Florida B 
Hood, S. C, 12224 Florida Avenue, Tampa, Florida B 
Houser, James G., Tech. Director, Technical and Research Staff, The Martin 

Company, P. O. Box 5837, Orlando, Florida P 
Hubbell, Dr. T. H., Univ. Michigan, Museum of Zoology, Ann Arbor, Mich- 
igan P 
Hubbs, Dr. Carl L., Scripps Institution of Oceanography, La Jolla, Califor- 
nia B 
Huddleston, Miss Carla J., Florida Southern College, Lakeland, Florida 



Membership List 171 

Hull, Robert W., Dept. Biological Sciences, Florida State Univ., Tallahassee, 

Florida 
Humphrey, Robert A., 430 East Kaley Ave., Orlando, Florida B 
Hunt, Burton P., Dept. Zoology, Univ. Miami, Coral Gables 46, Florida B 
Hunter, Dr. George W., Ill, Dept. Microbiology, Univ. Florida, Gainesville, 

Florida B 
Hutchman, Dr. J. E., P. O. Box 386, Lakeland, Florida P 
Hutton, Dr. Robert F., Division of Marine Fisheries, Dept. of Natural Re- 
sources, Carr's Landing, Wareham, Massachusetts B 
Ingle, Robert M., Florida State Board of Conservation, Tallahassee, Florida B 
Jerris, S. R., Roehr Products Co., Inc., P. O. Box 960, DeLand, Florida 
Johnson, Albert Smith, Chipola Junior College, Marianna, Florida B 
Johnson, Dr. Boyd W., Dept. Physics, Florida Presbyterian College, St. Peters- 
burg, Florida P 
Johnston, Dr. David W., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Jones, Dr. E. Ruffln, Jr., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Joyee, Edwin A., Jr., P. O. Box 136, St. Augustine, Florida B 
Joyner, Col. Houston C, 2025 Howard Dr., Winter Park, Florida 
Kaplan, Sherman R., 1680 Meridian Avenue, Miami Beach 39, Florida M 
Kelly, Robert D., 4011 San Rafael St., Tampa 9, Florida 
Kendall, Harry W., Physics Dept., Univ. South Florida, Tampa, Florida P 
Kilby, Dr. John D., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Kinser, B. M., P. O. Box 158, Eustis, Florida P 
Kinsey, P. E., 1647 Third Ave. N., Jacksonville Beach, Florida 
Knowles, Dr. Robert P., 2101 N. W. 25th Avenue, Miami, Florida M 
Koger, Dr. Marvin, Dept. Animal Sci., Univ. Florida, Gainesville, Florida B 
Kornegay, William F., 1408 N. W. 19th Ave., Ocala, Florida 
Krivanek, Dr. Jerome O., Univ. South Florida, Tampa, Florida B 
Kronsbein, Dr. John, Coll. Engineering, Univ. Florida, Gainesville, Florida P 
Lackey, Dr. James B., Box 497, Melrose, Florida B 

Laessle, Dr. Albert M., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Lakela, Dr. Olga, Univ. South Florida, Tampa, Florida B 
Lannutti, Joseph E., Dept. Physics, Florida State Univ., Tallahassee, Florida P 
Larson, Dr. Edward, Dept. Zoology, Univ. Miami, Coral Gables, Florida M 
Latina, Albert A., 311 A Science Bldg., Univ. South Florida, Tampa, Florida B 
Layne, Dr. James N., Dept. Conservation, Fernow Hall, Cornell Univ., Ithaca, 

New York B 
Leavitt, Dr. Benjamin B., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Leigh, Dr. W. Henry, Dept. Zoology. Univ. Miami, Coral Gables 46, Florida B 
Leto, Frank P., Jr., 4713 Leila Avenue, Tampa 11, Florida T 
Lewison, Leon Roy, Erwinstr. 72, Freiburg i. Breisgau, Germany 
Lillien, Irving, Dept. Chemistry, Univ. Miami, Coral Gables, Florida 
Lippincott, Sylvia C, 13762 74th Avenue North, Largo, Florida 
Lorz, Dr. Albert, 409 Newell Hall, Univ. Florida, Gainesville, Florida B 
Lovell, William V., Route 2, Box 18, Sanford, Florida P 
Luce, Samuel W., Route 1, Box 621, Lakeland, Florida S 
Lutz, Nancy E., Box 114, Mandarin, Florida 
Lyle, William R., Route 1, Box 148-B, Bartow, Florida B 



172 Quarterly Journal of the Florida Academy of Sciences 

McCloskey, Dr. James, 2905 Pinedale Ave., Lakeland, Florida 
McClure, J. S., 938 Bordeau Avenue W., Jacksonville 11, Florida P 
McCrone, Dr. John D., Florida Presbyterian College, St. Petersburg, Florida B 
McLean, Douglas E., 1281 N. W. 195th St., Miami 69, Florida 
Macgowan, Prof. Robert, Florida Southern College, Lakeland, Florida S 
Man, Dr. Eugene H., P. O. Box 8293, Univ. Miami, Coral Gables, Florida 
Marks, Dr. Meyer B., 1680 Meridian Beach, Miami Beach, Florida B 
Marshall, Dr. J. Stanley, School of Education, Florida State Univ., Tallahas- 
see, Florida S 
Mayer, John, 276-1 Corry Village, Gainesville, Florida 
Menzel, Dr. R. W., Oceanographic Institute, Florida State Univ., Tallahassee, 

Florida B 
Merrill, Mrs. Frank, 1504 River Hills Circle, Jacksonville, Florida P 
Miles, E. P., Computing Center, Florida State Univ., Tallahassee, Florida P 
Mills, Dr. Alfred P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida P 
Moe, Martin A., Jr., Board of Conservation, Marine Lab., P. O. Drawer F, 

St. Petersburg, Florida 
Monk, Dr. Carl D., Dept. Botany, Univ. Florida, Gainevsille, Florida B 
Montgomery, Joseph G., Manatee Junior College, Bradenton, Florida B 
Moody, Harold L., 545 N. Woodland, Winter Garden, Florida B 
Moore, Donald R., Marine Laboratory, 1 Rickenbacker Causeway, Va. Key, 

Miami 49, Florida 
Moore, McDonald, Hampton Junior College, Ocala, Florida 
Morgan, George B., Phelps Lab., E & I Bldg., Univ. Florida, Gainesville, 

Florida B 
Morris, David M., Dept. Anatomy, Univ. Miami, Coral Gables, Florida B 
Morton, Richard K., 2827 Holly Point Dr., Jacksonville 11, Florida S 
Mulson, Joseph F., Dept. Physics, Rollins College, Winter Park, Florida P 
Munn, Dr. Lottie, 641 Lake Avenue, Maitland, Florida P 
Murray, Mary Ruth, 1326 S. W. 1st St., Miami, Florida S 
Nation, Dr. James L., 537 N. W. 35th Terrace, Gainesville, Florida 
Neill, W. T., 122 Homecrest Rd., New Port Riehey, Florida B 
Nelson, Dr. Gid E., Jr., Univ. South Florida, Tampa, Florida B 
Nelson, Howard C, P. O. Box 162, Gainesville, Florida 
Noble, Dr. Nancy L., 1550 N. W. 10th Ave., Miami 36, Florida 
Norman, Mrs. Nelle B., 2731 Pine Summit Drive E., Jacksonville 11, Florida 
Ober, Lewis D., 1235 N. E. 204th St., N. Miami Beach 62, Florida B 
O'Brien, Dr. Aaron H., Dept. Biology, Stetson Univ., DeLand, Florida B 
Odom, Clifton T., 2301 S. 8th Ave., Arcadia, California B 
Olle, Miss Esther, 1310 Bryn Mawr, Chicago 40, Illinois B 
Olson, Dr. James Allen, Dept. Biochemistry, J. Hillis Miller Health Center, 

Univ. Florida, Gainesville, Florida B 
Orgell, Wallace H., Dept. Biological Sciences, Florida Atlantic Univ., Boca 

Raton, Florida B 
Orr, Robert K., 757 S. Johnson St., Lakeland, Florida 
Owens, E. G., Pensacola Junior College, Pensacola, Florida P 
Padgett, Herbert R., Jacksonville Univ., Jacksonville 11, Florida S 



Membership List 173 

Paff, Dr. George H., Dept. Anatomy, Univ. Miami School of Medicine, Coral 

Gables 34, Florida M 
Palmer, Dr. A. Z., Dept. Animal Husbandry and Nutrition, Univ. Florida, 

Gainesville, Florida B 
Park, Dr. Mary Cathryne, 450 Norwood St., Merritt Island, Florida S 
Patterson, Robert F., 233 E. 43rd St., Hialeah, Florida B 
Pearsall, Leigh M., Melrose, Florida S 

Pearson, Dr. Jay F. W., 515 Ingraham Bldg., Miami, Florida B 
Penner, Dr. Lawrence R., Dept. Zoology, Univ. Connecticut, Storrs, Connecti- 
cut B 
Perry, Rachel, 237 North Boulevard, DeLand, Florida B 

Phillips, Ronald C, Dept. Botany, Seattle Pacific College, Seattle 99, Wash- 
ington B 
Phipps, Dr. Cecil G., Box 181 A, Tennessee Tech, Cookeville, Tennessee P 
Pierce, Dr. E. Lowe, Dept. Biology, Univ. Florida, Gainesville, Florida B 
Pierson, William H., 1831 N. W. 10th Ave., Gainesville, Florida S 
Pirkle, Dr. E. C, 611 N. W. 35th St., Gainesville, Florida P 
Plendl, Dr. Hans S., Dept. Physics, Florida State Univ., Tallahassee, Florida P 
Plice, Dr. Max J., 324 River St., Palatka, Florida B 

Plyler, Earle K., Dept. Physics, Florida State Univ., Tallahassee, Florida P 
Poitras, Dr. Adrian W., Dept. Biology, Dade County Junior College, Miami, 

Florida B 
Polskin, Dr. Louis J., 1401 S. Florida Ave., Lakeland, Florida M 
Prichard, Elmer C, 605 N. Amelia, DeLand, Florida B 
Provost, Dr. Maurice W., Box 308, Vero Beach, Florida B 
Puryear, Dr. R. W., Florida Normal and Industrial Memorial College, St. 

Augustine, Florida S 
Rabkin, Dr. Samuel, 511 Sylvan Drive, Winter Park, Florida B 
Randel, William, Dept. English, Florida State Univ., Tallahassee, Florida S 
Rappenecker, Dr. Caspar, Dept. Geology, Univ. Florida, Gainesville, Florida P 
Ray, Dr. Francis E., P. 401, J. Hillis Miller Health Center, Univ. Florida, 

Gainesville, Florida B 
Ray, Dr. James D., Jr., Univ. South Florida, Tampa, Florida B 
Reid, Dr. George K., Dept. Biology, Florida Presbyterian College, St. Peters- 
burg, Florida B 
Reinsch, Dr. B. P., Florida Southern College, Lakeland, Florida P 
Reitz, Dr. J. Wayne, President, Univ. Florida, Gainesville, Florida S 
Reynolds, Dr. J. Paul, Dept. Zoology, Florida State Univ., Tallahassee, Flor- 
ida B 
Rich, Dr. Earl R., Dept. Zoology, Univ. Miami, Coral Gables 46, Florida B 
Rick, Prof. Henry, 1272 N. W. 172nd Terrace, Miami, Florida 
Rinckey, Gordon R., 1225— 59th St. S„ Gulfport, Florida 
Rivas, Luis Rene, Dept. Zoology, Univ. Miami, Coral Gables, Florida B 
Roberts, Dr. Leonidas H., Benton 202, Univ. Florida, Gainesville, Florida P 
Robins, Dr. C. Richard, Institute Marine Sci., Univ. Miami, 1 Rickenbacker 

Causeway, Va. Key, Miami 49, Florida B 
Rockwell, Leo L., 602 Charles, Lakeland, Florida S 
Ross, Arnold, P. O. Box 13351, Univ. Station, Gainesville, Florida P 



174 Quarterly Journal of the Florida Academy of Sciences 

Ross, Dr. John S., Dept. Physics, Rollins College, Winter Park, Florida P 
Roth, Eric M., Palm Lane Trailer Park, 13804 Nebraska Ave., Tampa, Florida 
Rowand, Tom, Route 3, Box 5A8, Lake City, Florida 
Sachs, Dr. K. Norman, Jr., U. S. Geological Survey, E-214 U. S. National 

Museum, Washington 25, D. C. P 
Sagawa, Dr. Yoneo, Dept. Botany, Univ. Florida, Gainesville, Florida B 
Sandstrom, Carl J., Dept. Biology, Rollins College, Winter Park, Florida B 
Saslow, Milton S., 4250 W. Flagler St., Miami 44, Florida M 
Saute, George, Dept. Mathematics, Rollins College, Winter Park, Florida P 
Sawyer, Dr. Earl M., Dept. Physics, Univ. Florida, Gainesville, Florida P 
Scheer, Edward W., Jr., 62 Forbes Rd., Milton 86, Massachusetts B 
Schneider, George H., 2292 S. W. 36th Ave., Miami 45, Florida 
Shultz, Dr. Harry P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida P 
Schwartz, Guenter, Dept. Physics, Florida State Univ., Tallahassee, Florida P 
Scolaro, Reginald J., Box 12645, University Sta., Gainesville, Florida P 
Scott, Bruce Von G., 6201 Chapman Field Dr., Miami 56, Florida P 
Scruggs, R. M., North Florida Junior College, Madison, Florida 
Seaman, Dr. Irvin, 470 Biltmore Way, Coral Gables, Florida M 
Shah, N. S., Dept. Pharmacology, Univ. Florida, Gainesville, Florida 
Sherman, Dr. H. B., 410 Howry Ave., DeLand, Florida B 
Shirley, Dr. Ray L., Nutrition Lab., Univ. Florida, Gainesville, Florida B 
Shor, Dr. Bernice C, Rollins College, Winter Park, Florida B 
Shuster, Dr. Carl N., 2035— 26th Ave. N., St. Petersburg 13, Florida P 
Shuttleworth, Dr. F. S., Div. Science and Math., Lake Sumter Junior College, 

Leesburg, Florida B 
Sickels, Dr. Jackson P., 541 San Esteban Ave., Coral Gables 46, Florida P 
Simon, Dr. Joseph L., Natural Science Div., Univ. South Florida, Tampa, 

Florida 
Simons, Dr. Joseph H., 1122 S. W. 11th Ave., Gainesville, Florida P 
Sims, Harold W., Jr., 120 Mt. Curve Ave., N. E., St. Petersburg, Florida 
Singletary, Mary L., Box 254, Kissimmee, Florida B 
Six, Dr. N. F., Jr., Scientific Research Laboratories, Brown Engineering Co., 

P. O. Drawer 917, Huntsville, Alabama P 
Slack, Dr. Francis, P. O. Box 706, Hobe Sound, Florida M 
Sleight, Dr. Virgil G., Dept. Geology, Univ. Miami, Coral Gables 34, Florida P 
Smith, Dr. Alex. G., Dept. Physics and Astronomy, Univ. Florida, Gaines- 
ville, Florida P 
Smith, Earl D., 2309 Coventry Ave., Lakeland, Florida P 
Smith, Dr. Frederick B., Dept. Soils, Univ. Florida, Gainesville, Florida B 
Smith, Marshall E., 418 W. Piatt St., Tampa 6, Florida M 
Smith, Cmdr. Nathan L., 631 N. W. 34th Dr., Gainesville, Florida P 
Smith, F. G. Walton, Univ. Miami, Marine Laboratories, Coral Gables, Flor- 
ida B 
Smith, Riley S., Jr., Rollins College, Winter Park, Florida 
Snyder, Carl H., Dept. Chemistry, Univ. Miami, Coral Gables 46, Florida P 
Snyder, Dr. Clifford Charles, 214 Alhambra Circle, Coral Gables, Florida M 
Sokoloff, Dr. Boris Th., Dept. Biology, Florida Southern College, Lakeland, 
Florida B 



Membership List 175 

Soule, Dr. Mortimer J., Jr., 2247 N. W. 11th Ave., Gainesville, Florida B 
South, Dr. D. E., Florida Presbyterian College, St. Petersburg, Florida P 
Spencer, Mrs. Marion, 215 Fox Place, Port Orange, Florida P 
Springer, Stewart, Bureau of Commercial Fisheries, North Rotunda, Museum 

Bldg., Stanford Univ., Stanford, California B 
Starke, Raymond R., 3760 Bayou Blvd., Pensacola, Florida 
Stevens, Miss Marion, 207 Hibiscus Dr., Miami Springs, Florida B 
Stewart, Mrs. Violet N., 439 Harbor Dr. S., Indian Rocks Beach, Florida 
Stormont, James C, 7340 S. W. 63rd Ave., Miami, Florida 
Stubbs, Sidney A., P. O. Box 52297, Houston, Texas 
Swann, Maurice E., 3101 W. 13th St., Panama City, Florida P 
Swanson, Dr. D. C, Dept. Physics, Univ. Florida, Gainesville, Florida P 
Swindell, David E., Jr., 905 E. Park Ave., Tallahassee, Florida B 
Sykes, James E., Biological Station, 75 — 33rd Ave., St. Petersburg Beach, 

Florida 
Tagatz, Marlin E., 210 A Pringle Circle, Green Cove Springs, Florida 
Tanner, W. Lee, Box 38, Lake Panasoffkee, Florida 

Tanner, William F., Dept. Geology, Florida State Univ., Tallahassee, Florida P 
Taylor, John L., 527 New York Ave., Dunedin, Florida 
Teas, Dr. Howard J., Program Director for Metabolic Biology, National Science 

Foundation, Washington 25, D. C. B 
Tebeau, Dr. Carl P., Dept. Chemistry, Univ. Miami, Coral Gables, Florida P 
Tebeau, C. W., Univ. Miami, 307 Aledo Ave., Coral Gables 46, Florida S 
Thomas, Dr. Dan A., Dean of the Faculty, Jacksonville Univ., Jacksonville 11, 

Florida P 
Thomas, Lowell P., The Marine Laboratory, 1 Rickenbacker Causeway, Va. 

Key, Miami 49, Florida B 
Thompson, Dr. B. D., 600 N. W. 36th St., Gainesville, Florida B 
Thompson, Dr. Claude E., 3321 Cesery Blvd., Jacksonville 11, Florida S 
Tinner, J. C, Morgan State College, Baltimore 18, Maryland P 
Tissot, Dr. A. N., Agricultural Exp. Sta., Univ. Florida, Gainesville, Florida B 
Topp, Robert, Florida Board of Conservation, Marine Lab., P. O. Drawer F, 

St. Petersburg, Florida 
Totten, Henry R., Dept. Botany, Univ. North Carolina, Box 247, Chapel Hill, 

North Carolina B 
Toulmin, Dr. Lymon D., Dept. Geology, Florida State Univ., Tallahassee, 

Florida P 
Truitt, John O., 808 Almeria Ave., Coral Gables 34, Florida 
Van Cleef, Alice, Glenwood, Florida P 

Van Vleck, David B., Dept. Zoology, Univ. Miami, Coral Gables, Florida B 
Vernon, Dr. Robert O., Box 631, Tallahassee, Florida P 
Vestal, Dr. Paul A., Dept. Botany, Rollins College, Winter Park, Florida B 
Voss, Dr. Gilbert L., Marine Laboratory, Univ. Miami, Coral Gables, Florida B 
Waldorf, Gray W., Pensacola Junior College, Pensacola, Florida B 
Wallace, Dr. H. K., Dept. Biology, Univ. Florida, Gainesville, Florida B 
Ward, Dr. Daniel B., 733 S. W. 27th St., Gainesville, Florida B 
Warnick, Dr. Alvin C, McCarty Hall 248, Univ. Florida, Gainesville, Flor- 
ida B 



176 Quarterly Journal of the Florida Academy of Sciences 

Warren, Cecil R., Crandon Park Zoo, 4000 Crandon Blvd., Miami, Florida B 
Watkins, Dr. Marshall O., 1115 N. E. 3rd St., Gainesville, Florida B 
Weber, Dr. George F., 406 Horticulture Bldg., Univ. Florida, Gainesville, 

Florida B 
Webster, W. C., Box 211, Gonzalez, Florida 
Wegner, W. E., 3100 Highland Rd., Baton Rouge, Louisiana 
Weigel, Dr. Robert D., Dept. Biology, Illinois State Univ., Normal, Illinois B 
Weisbord, Norman E., Dept. Geology, Florida State Univ., Tallahassee, Flor- 
ida P 
Weise, Gilbert N., 8601 Emerald Isle Circle N., Jacksonville 16, Florida 
Weiser, Dr. Josejf, Florida Memorial College, St. Augustine, Florida 
Wellman, Wayne E., 967 S. W. 5th St., Miami 36, Florida P and S 
Wells, Harry W., Dept. Biological Sciences, Florida State Univ., Tallahassee, 

Florida B 
West, Mrs. Felicia E., 6231 Spring Forest Circle, Jacksonville 16, Florida 
Westfall, Dr. Minter J., Jr., Dept. Biology, Univ. Florida, Gainesville, Flor- 
ida B 
Williams, Dr. James H., Rt. 1, Box 312, Avon Park, Florida S 
Williams, Louise lone, Lakeland High School, Lakeland, Florida B 
Williams, Lovett E., Jr., Game and Fresh Water Fish Comm., P. O. Box 908, 

Lake City, Florida B 
Williams, Dr. Robert H., Univ. Miami, P. O. Box 8233, Coral Gables 46, Flor- 
ida B 
Wilson, Druid, Room 404, U. S. National Museum, Washington 25, D. C. B 
Wilson, Dr. John L., 495 Savannah State College, Savannah, Georgia P 
Wingfield, E. Burwell, Virginia Polytechnic Institute, Box 6216, Va. Tech 

Station A, Blacksburg, Virginia 
Winthrop, Henry, Univ. South Florida, Tampa, Florida S 
Woodburn, Kenneth D., Fla. State Board of Conservation, Marine Lab., P. O. 

Drawer F, St. Petersburg, Florida 
Wolfenbarger, Dr. D. O., Sub.-Trop. Exp. Sta., Route 2, Box 508, Homestead, 

Florida 
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Florida B 
Wright, Mrs. Louise B., Chapman High School, Apalachicola, Florida P 
Wright, Dr. Peter C, Univ. South Florida, Tampa, Florida 
Wright, Shirley Jean, P. O. Box 4552, Miami Beach, Florida 
Yaffa, Harold, Dept. Biological Sciences, Florida State Univ., Tallahassee, 

Florida B 
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ida B 
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Young, Dr. Frank N., Dept. Biology, Univ. Indiana, Bloomington, Indiana B 
Zinner, Dr. Doran D., 2017 Alhambra Circle, Coral Gables, Florida B 
Zinober, Dr. M. R., P. O. Box 284, Live Oak, Florida 



FLORIDA ACADEMY of SCIENCES 

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for 1964 

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FLORIDA ACADEMY of SCIENCES 
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OFFICERS FOR 1964 

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WW 3 

/'Cfb$ Quarterly Journal 

of the 

Florida Academy of Sciences 



Vol. 27 



September, 1964 



No. 3 



CONTENTS 



Surface calcium-chlorinity relationships 
A new name for Fulica minor Shufeldt 



Billie Z. May 177 
Pierce Brodkorb 186 



New Eocene decapods from Florida 

Arnold Ross, Jackson E. Lewis, and R. J. Scolaro 187 



Survival potential of piranhas in Florida 
New subspecies of Leiocephalus from Cuba 
Notes on fossil turkeys 
Osteology of gallinaceous birds 



Martin A. Moe, Jr. 197 

Albert Schwartz 211 

Pierce Brodkorb 223 

/. Alan Holman 230 




Mailed November 4, 1964 



Quarterly Journal of the Florida Academy of Sciences 
Editor: Pierce Brodkorb 



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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY OF SCIENCES 



Vol. 27 September, 1964 No. 3 

SURFACE CALCIUM-CHLORINITY RELATIONSHIPS 
BiLLiE Z. May 

Concentrations of calcium ions directly affect the existence 
and distribution of aquatic organisms (De Sousa, 1954; Borg, 1931). 
This element plays a significant role in cellular synthesis of phyto- 
plankton (Robertson, 1941). Calcium studies are a prominent part 
of numerous ecological surveys of marine, brackish, and fresh- 
water ecosystems (Remane and Schlieper, 1958; Vinogradov 1953). 

Literature pertaining to calcium in the waters of the west coast 
of Florida is scarce. The only available data are those of Drago- 
vich, Finucane, Kelly, and May (1963). Data forming the base for 
this research were collected as part of the red-tide and estuarine 
ecological investigations conducted by the Bureau of Commercial 
Fisheries. The objective of this study was to develop formulae 
for the rapid determination of calcium in surface waters of Tampa 
Bay and the adjacent neritic area. 

Materials and Methods 

The investigation of the calcium content of surface waters was 
initiated in April 1960 at stations already being systematically sur- 
veyed for other chemical, physical, and biological parameters (Dra- 
govich et at., 1963). Duration of the sampling period was 16 
months, continuing through July 1961. No samples were taken 
in April 1961. Surface samples were collected monthly at all 
stations (fig. 1). At stations 1 through 20, sampling occurred with- 
in one hour of high tide. Stations 21 through 25 were sampled 
regardless of tidal stage. 

Water samples for calcium and chlorinity analysis were col- 
lected with modified Van Dorn sampling bottles (Van Dorn, 1957). 



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The aliquots for calcium were dispensed into 250 ml reagent bot- 
tles and capped with ground glass stoppers. Samples for chlorin- 
ity measurements were dispensed into 4-ounce prescription bottles 
and closed with standard lined-plastic caps. 

The large number of water samples to be analyzed in the lab- 
oratory necessitated selection of a fast, dependable method for de- 
termining calcium content in sea water. The classic method is to 
precipitate calcium as the oxalate and ignite the precipitate to the 
oxide (Thompson and Wright, 1930; Matida, 1950). This method 
was considered too time-consuming to lend itself well to estuarine 
investigations. The same objection applied to the calcium oxalate 
precipitation method which involves dissolution of the oxalate in 
sulfuric acid and titration of the oxalic acid with potassium per- 
manganate (Kirk and Moberg, 1933; Gripenberg, 1937). Direct ti- 
tration of sea water with disodium ethylene-diaminetetracetate 
(EDTA) is a method which embodies rapidity, east of manipula- 
tion, and negligible error. This method was selected for the cal- 
cium determinations in the present study. The error resulting from 
the complexometric titration is negligible because of the weak iron 
content of sea water (less than 1 ppm) and trace amounts of alumi- 
num and other elements. 

In the EDTA titration of sea water for calcium, the procedure 
was as follows: An aliquot of 100 ml was adjusted with sodium 
hydroxide to a pH slightly greater than 12.0. A suitable amount 
(0.6 ml) of a 0.1 per cent solution of murexide (ammonium purpu- 
rate) was added as an indicator and the solution titrated with 0.1 
molar EDTA until the indicator turned blue-violet. Other ions 
present do not interfere in this procedure. The Ca +2 concentration 
was then calculated in milligram-atoms per liter, assuming that, 
stoichiometrically, 1 ml of the reagent was equivalent to 4.008 mg. 
of calcium. Calcium values in this study were reported in milli- 
grams per kilogram of sea water. 

Chlorinity measurements were made according to a modified 
version of the Knudsen method (Oxner, 1920) with a standard sil- 
ver nitrate titration. Replacement of the Knudsen burette by an 
automatic Schellbach burette facilitated the evaluation of low level 
chlorinities. Other modifications in the equipment and procedure 
are described as follows: A 15 ml portion of the sample was with- 
drawn with an automatic pipette and discharged into a 100 ml Ber- 
zelius beaker. The aliquot was placed on a magnetic stirrer and 



180 



Quarterly Journal of the Florida Academy of Sciences 




May: Catcium-Chlorinity Relationships 181 

an adequate quantity (1 ml) of an 8 per cent solution of potassium 
chromate added as an indicator. The aliquot was titrated with sil- 
ver nitrate solution subsequent to the adjustment of the titer so as 
to be approximately equivalent in double milliliters (2 ml per 
1 o/oo CI) to the chlorinity of LA.P.O. standard sea water (19.376 
o/oo CI). The end point was taken as the change in color from 
yellow to faint red which persisted for 30 seconds. The chlorinity 
was corrected by use of correction values in Knudsen's Hydro- 
graphical Tables (Knudsen, 1901). 

Results 

Stations 1 through 20 were considered as inshore area, while 
stations 21 through 25 were designated as offshore (fig. 1). This 
classification is based upon calcium and salinity variations in both 
areas. In inshore and offshore areas the salinity varied 138 per 
cent and 2 per cent, respectively. The calcium variation was of 
similar magnitude in that the inshore variation was 129 per cent 
and the offshore 13 per cent. 

The means of the fifteen calcium values for each station were 
plotted against the means of the chlorinities for the inshore area 
(fig. 2). The analogous values for offshore water were treated sim- 
ilarly (fig. 3). A straight line relationship was found to exist for 
the range of values encountered. 

An equation for each straight line was computed by the method 
of least squares. Individual values were used in the calculations 
rather than the means shown in the figures. In the inshore area 
chlorinity and calcium values varied from 7.73 to 19.81 o/oo and 
from 173.2 to 426.0 mg/kg, respectively. The range of chlorinity 
and calcium in the offshore area was 18.06 to 20.23 o/oo and 382.8 
to 434.1 mg/kg, respectively. For inshore waters the derived 
formula (equation 1) was: 

Ca (g/kg) = 0.0154 + 0.0207 CI (o/oo). 

The formula (equation 2) for the offshore area was: 

Ca (g/kg) = 0.0004 + 0.02137 CI (o/oo). 

Both formulae were tested for accuracy by comparing titrated cal- 
cium values with derived values from 52 sea water samples. These 
samples were collected from locations originating at the mouths 



182 



Quarterly Journal of the Florida Academy of Sciences 




bo 



May: Calcium-Chlorinity Relationships 



18' 



TABLE 1 

Standard and average deviation between calculated 
and analytical values of calcium 









A. Inshore Formula 










(m 


g/kg) 






(mg/kg) 




Sample 


Anal. 


Formula 


Diff. 


Sample 
21 


Anal. 
384.5 


Formula 
382.0 


Diff. 


1 


246.7 


240.6 


-6.1 


-2.5 


2 


249.9 


241.7 


—8.2 


22 


379.6 


377.6 


-2.0 


3 


251.1 


242.7 


-8.4 


23 


361.2 


354.7 


-6.5 


4 


251.5 


244.8 


-6.7 


24 


359.0 


349.7 


—9.3 


5 


251.8 


246.0 


-5.8 


25 


392.2 


388.8 


-3.4 


6 


250.3 


243.9 


-6.4 


26 


375.1 


383.6 


+8.5 


7 


252.6 


245.4 


-7.2 


27 


319.3 


313.7 


—5.6 


8 


251.5 


244.8 


-6.7 


28 


387.9 


387.6 


-0.3 


9 


248.3 


244.8 


—3.5 


29 


345.1 


340.8 


-4.3 


10 


286,3 


288.4 


+2.1 


30 


288.2 


273.7 


-14.5 


11 


253.3 


251.0 


—2.3 


31 


383.8 


381.8 


-2.0 


12 


245.2 


243.7 


—1.5 


32 


383.4 


381.6 


—1.8 


13 


224.2 


217.0 


—7.2 


33 


392.5 


391.1 


-1.4 


14 


203.5 


196.1 


—7.4 


34 


393.3 


390.5 


-2.8 


15 


322.2 


332.5 


+ 10.3 


35 


385.3 


403.9 


+ 18.6 


16 


285.9 


299.0 


+ 13.1 


36 


362.4 


373.3 


+ 10.9 


17 


302.4 


307.9 


-5.5 


37 


322.0 


327.6 


+4.7 


18 


280.8 


267.3 


— 13.5 


38 


283.2 


285.7 


+2.5 


19 


290.2 


271.0 


—19.2 


39 


389.5 


398.7 


-9.2 


20 


382.4 


378.3 


-4.1 


40 


368.0 


378.2 


— 10.2 



Average Deviation = 6.7 mg/kg or 2.1% error 
Standard Deviation = 8.0 mg/kg 



B. Offshore Formula 



1 


398.6 


399.4 


+0.8 


2 


398.6 


396.5 


-2.1 


3 


401.9 


399.4 


-2.5 


4 


407.0 


406.0 


—1.0 


5 


416.2 


419.9 


+3.7 


6 


403.9 


404.8 


+0.9 



7 


395.0 


413.3 


+ 18.3 


8 


408.5 


419.9 


+ 11.4 


9 


406.2 


404.5 


+ 1.7 


10 


402.7 


404.7 


—2.0 


11 


412.2 


414.1 


-1.9 


12 


406.9 


409.2 


-2.3 



Average Deviation = 4.0 mg/kg or 1.0% error 
Standard Deviation = 6.9 mg/kg 



184 Quarterly Journal of the Florida Academy of Sciences 

of Old Tampa and Hillsborough Bays to a distance of 40 miles off- 
shore. The calcium levels in these samples were determined an- 
alytically by titration and also by applying the applicable Ca/Cl 
formula, subsequent to titration of the chlorinity. The average 
deviation of calculated values from analytical values for the inshore 
area was 6.7 mg/kg (table 1). This represented an average error 
of 2.1 per cent in that area. The standard deviation amounted to 
8.0 mg/kg. For the offshore area, the average deviation was 4.0 
mg kg or 1.0 per cent error. The standard deviation was 6.9 
mg/kg. 

Discussion 

Noticeable differences in the Ca/Cl ratio have been observed 
in various marine areas of the world (Sverdrup, Johnson, and Flem- 
ing, 1942). No agreement was found between our Ca/Cl ratios 
observed in the inshore area and those of oceanic waters (Kirk and 
Moberg, 1933; Thompson and Wright, 1930; Chow and Thomp- 
son, 1955; Carpenter, 1957). 

Equating CI = in equation 1, an intercept value of 0.0154 
is found which indicates that the offshore waters entering Tampa 
Bay are diluted with fresh water containing 15.4 milligrams of 
calcium per kilogram of water. The surface waters of the immedi- 
ate offshore area approach a direct ratio closely since in this in- 
stance the intercept value in equation 2 is 0.0004. 

By utilizing these two formulae for the areas specified, calcium 
values may now be determined without recourse to chemical anal- 
ysis. The only analytical requirement is an accurate measurement 
of chlorinity which can be obtained with a conductivity cell. This 
procedure for the determination of calcium presents a rapid and 
convenient method. The reduction in laboratory expenditures and 
time is of practical significance. 

Literature Cited 

Borg, F. 1931. On some species of Membranipora. Arkiv. Zool. Stock- 
holm, vol. 22. 

Carpenter, J. H. 1957. The determination of calcium in natural waters. 
Limnol. Oceanogr., vol. 2, pp. 271-280. 

Chow, J. J., and T. G. Thompson. 1955. Flame photometric determination 
of calcium in sea water and marine organisms. Anal. Chem., vol. 27, 
pp. 910-913. 



May: Calcium-Chlorinity Relationships 185 

Dragovich, A., J. H. Finucane, J. A. Kelly, Jr., and B. Z. May. 1963. 
Counts of red tide organisms, Gymnodinium breve, and associated 
oceanographic data from Florida west coast, 1960-1961. U. S. Fish 
and Wildlife Serv., Spec. Sci. Rept., Fish. no. 445, 40 pp. 

DeSousa, Arthur. 1954. La determination rapide du calcium et du mag- 
nesium dans 1'eau de mer. Anal. Chim. Acta, vol. II, pp. 221-224. 

Gripenberg, Stina. 1937. A simplified method for the determination of cal- 
cium in sea water. Jour. Cons. Internat. Explor. Mer, vol. 12, pp. 
284-292. 

Kirk, P. L., and E. G. Moberg. 1933. Micro-determination of calcium in 
sea water. Ind. Eng. Chem. Anal. Ed., vol. 5, pp. 95-97. 

Knudsen, M. 1901. Hydrographical tables. Tutein & Koch, Copenhagen. 
63 pp. 

Matida, Yoshihdro. 1950. On the chemical composition of sea water in the 
Tokyo Bay. Jour. Ocean. Soc. Japan, vol. 5, pp. 105-110. 

Oxner, M. 1920. Manuel pratique de l'analyses de l'eau de mer. I. Chloru- 
ration par la methode de Knudsen. Bull, de la Comm. Internat. pour 
l'Explor. Scient. de la Mer Mediterranee, vol. 3. 

Remane, A., and C. Schlieper. 1958. Die Biologie des Brackwassers. Vol. 
22, E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, 348 pp. 

Robertson, J. D. 1941. The function and metabolism of calcium in the in- 
vertebrate. Biol. Rev., vol. 16, pp. 106-133. 

Sverdrup, H. U., M. W. Johnson, and R. H. Fleming. 1942. The Oceans. 
Prentice-Hall, Englewood Cliffs, N. J., 1087 pp. 

Thompson, T. G., and C. C. Wright. 1930. Ionic ratios of the waters of 
the North Pacific Ocean. Jour. Amer. Chem. Soc, vol. 52, pp. 915- 
921. 

Van Dorn, W. G. 1957. Large-volume water sampler. Trans. Amer. 
Geophys. Union, vol. 37, pp. 682-684. 

Vinogradov, A. P. 1953. The elementary chemical composition of marine 
organisms. Sears Foundation for Marine Research, New Haven, pp. 
647. 

Bureau of Commercial Fisheries Biological Station, St. Peters- 
burg Beach, Florida. Contribution No. 13. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



A NEW NAME FOR FULICA MINOR SHUFELDT 
Pierce Brodkorb 

The Pleistocene coot of Fossil Lake, Oregon, was briefly de- 
scribed and later figured by Shufeldt (1891, pp. 819, 820; 1892, 
p. 412, pi. 17, fig. 32) under the name Fulica minor, the type being 
a humerus, no. 3480 in the American Museum of Natural History. 
Howard (1946, p. 182) studied 161 bones from Fossil Lake and 
reduced the form to a temporal subspecies of the living American 
coot, as Fulica americana minor Shufeldt. Wetmore (1956, p. 57) 
subsequently reinstated it to full specific rank. 

In any case the name is preoccupied by Fulica minor Brehm 
(1831, p. 711) for a South American species. For Fulica minor 
Shufeldt I therefore propose the new name Fulica shufeldti. 

Literature Cited 

Brehm, Christian Ludwig. 1831. Handbuch der Naturgeschichte aller 
Vogel Deutschlands. Bernh. Friedr. Voigt, Ilmenau, xxiv + 1088 pp., 
colored frontispiece, 46 colored pis. 

Howard, Hildegarde. 1946. A review of the Pleistocene birds of Fossil 
Lake, Oregon. Carnegie Instn. Washington Publ., no. 551, pp. 141- 
195, pis. 1-2. 

Shufeldt, R. W. 1891. Fossil birds from the Equus beds of Oregon. Amer. 
Naturalist, vol. 25, no. 297, pp. 818-821. 

. 1892. A study of the fossil avifauna of the Equus beds of the Ore- 
gon desert. Jour. Acad. Nat. Sci. Philadelphia, vol. 9, pp. 389-425, pis. 
15-17. 

Wetmore, Alexander. 1956. A check-list of the fossil and prehistoric birds 
of North America and the West Indies. Smithsonian Misc. Coll., vol. 
131, no. 5, pp. 1-105. 

Department of Biology, University of Florida, Gainesville, 
Florida. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



NEW EOCENE DECAPODS FROM FLORIDA 
Arnold Ross, Jackson E. Lewis, and R. J. Scolaro 

The decapod crustacean fauna of the Florida Eocene has re- 
ceived little attention since the studies by Rathbun (1929, 1935) 
and Roberts (1953). Recently, Ross and Scolaro (1964) described 
a new species of Calappilia from the Williston Member of the Ocala 
Limestone of late Eocene age. Further studies on the Florida 
fauna disclose two additional new species from the Williston, one 
of the genus Calappa and the second interpreted as belonging to 
an undescribed but related genus. 

The genus Calappa is well represented in sediments of middle 
and late Tertiary age (Fig. 1), but Eocene representatives of this 
group are virtually unknown. Predicated on the existing litera- 
ture, the authors infer that the Calappa complex arose near the end 
of the Eocene. The new genus probably arose from the same 
stock during the early or middle Eocene and presumably became 
extinct toward the close of this epoch. 

The morphological terminology employed throughout this 
study follows that proposed by Rathbun (1918) and modified sub- 
sequently by Holthuis (1958). 

Institutional abbreviations used are as follows: Paleontological 
Research Institution, Ithaca, P.R.I. ; Florida State Museum, F.S.M. 

All of the specimens here described and figured were collected 
by Dr. Harold K. Brooks from the bottom of a limestone sink in 
the Williston Member of the Ocala Limestone locally referred to as 
"Devil's Den." The sink is in the S.E. V*, Sec. 26, T. 12 S., R. 18 E., 
about two miles north and one mile west of Williston, Levy 
County, Florida. A stratigraphic section of the sink has been pre- 
sented by Floyd (1962). 

Family Calappidae Dana, 1852 

Subfamily Calappinae Alcock, 1896 

Genus Calappa Weber, 1795 

Calappa robertsi, new species 

Figs. 2a-d 

Diagnosis. Outer surface of the propodus is distinctly zoned, 
bearing tubercles and granules. The upper zone (3) bears 15 



188 Quarterly Journal of the Florida Academy of Sciences 



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Ross, Lewis, Scolaro: New Eocene Decapods 189 

pustulose tubercles arranged in four rows. Zone 1 is delimited 
from zone 3 by a sparsely granulated trough. The cristate super- 
ior margin of both propodites carries six prominent teeth. 

Description. The outer surface of the manus is divided into 
three distally widening zones which are slightly oblique to the 
inferior margin. The lowermost zone (1) bears numerous medium 
granules arranged in sub-parallel rows and is set off from the next 
higher, second zone by an indistinctly defined ridge. The ridge 
carries one minor and two major tubercles; the minor tubercle is 
proximal. On the major (right) propodus, the stout, lobular pro- 
jection bears the distal major tubercle on its superior slope. The 
tubercles are finely granular, creating a pustulose appearance. The 
second zone (2) occupies a shallow trough or sulcus and bears 
randomly oriented granules similar to, although less abundant 
than, those in zone 1. The uppermost zone (3) occupies approxi- 
mately two-thirds of the outer surface and bears 15 major and 
minor pustulose tubercles arranged in four irregular, sub-parallel 
rows set among randomly oriented medium granules. The first 
of these rows delimits zone 3 from zone 2 and consists of one 
minor and four major tubercles; the minor tubercle adjoins the 
proximal one. The other three rows, in ascending order, consist of 
three major tubercles, one minor and three major tubercles, and 
one minor and two major tubercles, respectively. The minor tu- 
bercle in row three adjoins the distal one. The minor tubercle 
in row four lies on the base of the first (distal) crestal tooth, 
while the associated major tubercles are situated on the surface 
of the manus, inferior to and between the third and fourth teeth 
and the fifth and sixth teeth. 

The distal extremity of the propodus bears a large, laterally 
compressed supra-dactylar projection, the margin of which is 
finely crenate. The cockscomb crowning the superior margin of 
both major and minor propodites bears six erect, pointed teeth. 
A seventh, low, two-topped tooth is situated on the supero-proximal 
slope. The articular sockets of the propodus, which correspond 
to processes on the carpus, are deep, and the processes are prom- 
inent. A pointed, triangular, lamellated tooth is situated at the 
infero-proximal extremity of the outer surface. The inferior mar- 
gin is almost straight and is widest at the infero-proximal extremity, 
where it is confluent with the lamellar tooth. The margin gradu- 
ally thins distally for about two-thirds of its overall length until 



190 Quarterly Journal of the Florida Academy of Sciences 




Fig. 2. Calappa robertsi, new species, a, b, external and internal views 
of left propodus, hole-type, P.R.I. No. 6064 (actual height 20.9 mm.); c, ex- 
ternal view of right propodus, paratype, P.R.I. No. 6065 (actual height 13.5 
mm.); d, external view of left propodus, paratype, F.S.M. No. 1334 (actual 
height 16.4 mm.). Aparnocondylus ocalanus, new genus and species, e, f, 
external and internal views of right propodus, holotype, F.S.M. No. 1338 
(actual height, 20.1 mm.). 



Ross, Lewis, Scolaro: New Eocene Decapods 191 

it begins to grade into the fixed finger, where it is flexed sharply 
downward and inward. Granulation is random over the broad 
portion of the margin, becoming resolved at the point of flexure 
into three parallel rows, which continue on the finger. The inferior 
inner margin of the propodus is crenate, marked by a slightly ele- 
vated row of closely spaced, bead-like granules. The inner sur- 
face of the manus is very finely granulated, the granules becoming 
larger in a concentration adjacent to the inter-digital depression, 
where they are suggestive of a stridulating organ. An obscure 
line of widely spaced, larger granules emerges from the center 
of the concentration and extends diagonally across the upper inner 
surface of the fixed finger to meet the primary teeth distally. Traces 
of five primary teeth remain on the superior margin of the finger, 
extending from the inferior margin of the dactylar orifice to the 
point where the extremity of the finger has been destroyed. A low, 
flat, obliquely inclined ridge, extending along the inner surface of 
the finger adjacent to the inferior margin, emanates from a small 
node midway between the point of flexure and the extremity. 

Granules on the inner and outer surfaces of the fixed finger and 
along the distal inferior margin of the propodus are punctate, 
probably having facilitated passage of tufts of sensory hairs during 
life. 

The upper outer surface of the carpus bears medium granules 
and at least one row of pustulose tubercles sub-parallel to the 
superior margin; the inner surface bears fine granules only. The 
superior margin is arcuate and crenate, bearing an elevated row 
of closely spaced, small granules which become progressively larger 
distally. The supero-distal extremity terminates in a short, sharp 
spine. The articular process at the superior margin of the propodal 
orifice is elongate and prominent. The outer edge of the orifice 
bears a poorly defined row of small, bead-like granules. 

The supero-distal external surface of the merus is coarsely gran- 
ular between the transverse, wing-like expansion and the distal 
margin. The upper portion of the expansion is depressed and 
flat. The expansion bears at least three broad, prominent, pustulose 
teeth which are inclined ventrally. Like the carpus, the superior 
margin and the outer edge of the carpal orifice are both finely 
crenate. 

Remarks. At the present time Calappa robertsi n. sp. is the 
only identifiable North American Eocene representative of the 



192 Quarterly Journal of the Florida Academy of Sciences 

genus Calappa. Of the Recent species examined, only two show 
close affinities with this new species. Calappa gallus (Herbst) dif- 
fers in the less distinct zonation of the manus; no zone 2 sulcus; 
fewer teeth comprising the cockscomb, which is more highly ele- 
vated; and the greatly reduced infero-proximal spine and supra- 
dactylar projection. The propodus of C. angusta Milne-Edwards is 
much more tuberculate; zone 2 is not depressed and bears several 
tubercles in addition to granules; the inferior margin is sharp and 
crenate; and the cockscomb extends distally farther, relative to 
the supra-dactylar projection. 

Measurements of Holotype. Overall length 24.9, height 20.9, 
sub-length 21.9, sub-height 7.4 mm. The overall length of the 
propodus, as defined here, is the distance along the inferior margin 
from the distal end of the fixed finger to the infero-proximal ex- 
tremity. The height is the greatest perpendicular distance between 
the superior and inferior margins. The distance between the in- 
ferior margin of the dactylar orifice and the inferior point of articu- 
lation with the carpus is referred to as the sub-length. The sub- 
height is the shortest distance between the inferior margin of the 
dactylar orifice and the inferior margin of the propodus. 

Measurements of Paratypes. The measurements of paratype 
F.S.M. No. 1334 (left propodus) are as follows: overall length 19.5, 
height 16.4, sub-length 17.3, sub-height 6.0 mm. The remaining 
specimens in the paratypic lot are fragments. All of the fragments 
appear to belong to individuals of the same size as, or smaller than, 
the specimen represented by the holotype. 

Type Depositories. The holotype and one paratype have been 
deposited in the Paleontological Research Institution collections, 
catalogue numbers 6064 (left propodus) and 6065 (right manus), 
respectively. The remaining paratypes are in the Florida State 
Museum collections at the University of Florida, catalogue num- 
bers 1334 (left propodus), 1335 (left manus; fragment), 1336 (left 
merus; fragment), and 1337 (right carpus; fragment). 

Distribution. One left propodus was collected by the senior 
author from a limerock quarry at the town of Haile, Sec. 13, T. 
9 S., R. 17 E., Alachua County, Florida; Williston Member, Ocala 
Limestone, Upper Eocene. 

Etymology. The authors take great pleasure in naming this 
new species in honor of Henry B. Roberts of the United States 
National Museum. 



Ross, Lewis, Scolaro: New Eocene Decapods 193 

Subfamily indet. 
Genus Aparnocondylus, new genus 

Definition. The calappid genus Aparnocondylus is closely re- 
lated to the genus Calappa Weber, 1795. The new genus may be 
easily distinguished from all genera in the subfamily Calappinae 
by the absence of a stout, lobular projection on the outer surface 
of the propodus in the region where the fixed finger joins the 
manus. The propodus is laterally compressed, arcuate in cross- 
section, and the superior margin of the manus forms a high den- 
tate crest. 

Etymology. From Greek, aparnos (denying utterly) and kondu- 
los (masculine, a knob or prominence). 

Type Species. Aparnocondylus ocalanus, new species. 

Remarks. Based on the limited material available, the authors 
do not feel justified in assigning the monotypic genus Aparnocon- 
dylus to a subfamily at this time, although the observed character- 
istics appear to warrant the erection of a new subfamily. That 
the asymmetry of propodites exhibited in all living genera of 
Calappinae had become established in the Eocene is confirmed 
by Calappa robertsi new species (see Fig. 2c). However, Aparno- 
condylus is interpreted as indicative of a more primitive evolution- 
ary condition. It is improbable that the type of A. ocalanus repre- 
sents a reversal of asymmetry associated with pathological mal- 
formation. 

Aparnocondylus ocalanus, new species 
Figs. 2e-f 

Diagnosis. The outer surface of the propodus has low tuber- 
cles and granules. Zone 2 is a non-depressed plain. Zone 3 
bears 25 tubercles; the zone is divisible into four rows of minor 
and major tubercles. The areas between rows three and four and 
row four and the crest are arcuate and devoid of tubercles. 

Description. The outer surface of the manus is divided into 
three major zones, which are oriented slightly oblique to the plane 
of the inferior margin. The lowermost zone (1) bears many large, 
closely spaced, randomly arranged granules, which also cover the 
outer surface of the fixed digit. Proximally, the zone is narrow, 
originating distal to the infero-proximal tooth but rapidly widening 
and terminating distally at the inferior margin of the dactylar ori- 
fice. Zone 1 is delimited from the next higher zone by a row of 



194 Quarterly Journal of the Florida Academy of Sciences 

five very finely granular tubercles. The proximal pair is composed 
of one minor and one, slightly inferior and distal, major tubercle; 
the distal triplet has one major tubercle between two minor tu- 
bercles. Zone 2 occupies a flat, narrow plain of uniform width 
and bears few small and fewer medium granules. The uppermost 
zone (3) includes a little over one-half of the outer surface and bears 
25 major and minor tubercles arranged in four irregular rows in a 
ground-mass of medium and small granules. The first of these rows 
sets off zone 3 from zone 2 and consists of four major and two 
minor tubercles. The minor distal tubercle is adjacent to the outer 
dactylar furrow; the other minor tubercle adjoins the proximal 
major one. Row two contains two groups of three tubercles, a 
proximal group of one major between two minors, and a medial 
group of three minor tubercles. Of the seven major tubercles in 
row three, the distal two are joined, one obliquely above the 
other, and a smaller major tubercle lies above and between two 
large ones in the central portion of the manus. Row four consists 
of one minor and five major tubercles arranged in three groups. 
The minor tubercle is proximal and is associated with two smaller 
major ones; a single major tubercle occurs on the inferior basal 
slope of the supra-dactylar projection; a medial pair of major tu- 
bercles lies between the other groups. While the first two rows 
are straight and parallel, the third curves slightly and the fourth 
is strongly arcuate, parallel to the superior margin of the manus. 
Rows one, two, and three are closely spaced, but row four is sep- 
arated from the underlying row and the overlying cockscomb by 
uniformly wide, arcuate, granulated areas. 

The superior margin of the propodus bears a cockscomb of 
seven teeth. The distal pair appear to have been partially fused 
and the proximal tooth is bifid. A prominent, laterally compressed, 
supra-dactylar projection marks the supero-distal extremity. The 
outer surfaces of the teeth and the supra-dactylar projection bear 
medium to small granules. 

The inferior margin of the propodus is broadly arcuate. The 
proximal portion occupies approximately two-thirds of the overall 
length of the propodus and is broad, flat, and weakly granulated; 
the distal one-third is tapered and rounded. Strong granules on the 
distal third form two parallel rows which continue along the in- 
ferior margin of the fixed digit. A third, weaker row, parallel to 
the other two, delimits the entire inferior margin from the inner 



Ross, Lewis, Scolaro: New Eocene Decapods 195 

surface of the manus. It is more strongly developed toward both 
extremities, and is terminated distally, immediately below the 
emergence of a salient ridge on the inner surface of the fixed finger. 
A broad, triangular, lamellar tooth rises from the infero-proximal 
extremity. The fixed digit curves sharply and obliquely inward . 

The articular processes on the proximal slope of the propodus, 
which correspond to depressed areas on the carpus, are triangular 
and bulbous. Each process bears a deep pit on its expanded ar- 
ticulating surface, probably accommodating a comparable projec- 
tion on the corresponding carpal process. 

The inner surface of the propodus is locally granular. Two rows 
of granules emerge from the top of the inner dactylar furrow, curv- 
ing a short distance obliquely downward and proximally. The 
superior row contains one small and four large granules; the proxi- 
mal large granule is inserted immediately above the small one. 
The inferior row consists of medium granules grading into a weak 
concentration of randomly oriented, small granules which occupies 
the distal one-half of a shallow trough. The trough is almost 
straight and extends from the inferior margin of the dactylar ori- 
fice, where it is grooved, to the inferior articular socket on the prox- 
imal slope. A weakly defined ridge, bearing widely spaced medium 
granules, originates on the inferior slope of the trough at the base 
of the fixed finger and extends distally along the axis of the digit. 
A broad, slightly elevated, arcuate platform curves across the 
manus, above the trough, from the supra-dactylar projection to 
the inferior margin of the carpal orifice. A horizontal row of three 
large, well separated granules is situated on the inferior slope of 
the platform, midway between the extremities. A broad, shallow, 
irregularly ovate depression occupies the distal area between the 
cockscomb and the arcuate platform. 

Granules on the outer surface of the fixed finger, as well as the 
ones on and near the inferior and distal margins of the propodus, 
are punctate, indicating the presence of heavy concentrations of 
sensory hairs during life. The larger granules on the inner surface 
are punctate also. 

Remarks. Although Aparnocondylus ocalanus bears in common 
with species of Calappa similar proportions, high cristate superior 
margin, and a tuberculate zoned surface, the absence of a lobular 
projection on the right propodus of A. ocalanus should serve to 
distinguish the new species. 



196 Quarterly Journal of the Florida Academy of Sciences 

Measurements of Holotype. Overall length 24.1 mm., height 
20.1 mm., sub-length 21.5 mm., sub-height 7.2 mm. 

Type Depository. The holotype has been placed in the Florida 
State Museum collections at the University of Florida, catalogue 
number 1338. 

Etymology. The specific name, ocalanus, reflects the fact that 
the species is from the Ocala Limestone. 

Acknowledgments 

The authors are grateful to Dr. Harold K. Brooks, Curator of 
Invertebrate Paleontology, Florida State Museum, for making 
available the fossil material. They are indebted also to Dr. Austin 
B. Williams of the University of North Carolina Institute of Fish- 
eries Research at Morehead City, who arranged the loan of Recent 
comparative specimens and critically read the manuscript. Henry 
B. Roberts of the United States National Museum critically read 
the species descriptions and provided additional Recent calappid 
crabs for comparison. 

Literature Cited 

Floyd, J. G. 1962. Stratigraphic distribution of Upper Eocene larger for- 
aminifera from north eastern Levy County, Florida. Unpubl. Univ. 
Florida master's thesis, pp. 1-55, pis. 1-3, text figs. 1-7. 

Holthuis, L. B. 1958. West Indian crabs of the genus Calappa, with a 
description of three new species. Studies Fauna Curacao, vol. 8, pp. 
148-186, figs. 28-54. 

Rathbun, Mary J. 1918. The grapsoid crabs of America. Bull. U. S. Nat. 
Mus., no. 97, pp. 1-461, text figs. 1-172, pis. 1-161. 

. 1929. A new crab from the Eocene of Florida. Proc. U. S. Nat. 

Mus., vol. 75, art. 15, pp. 1-4, pis. 1-3. 

. 1935. Fossil Crustacea of the Atlantic and Gulf Coastal Plain. 

Geol. Soc. Amer., Spec. Paper no. 2, pp. 1-160, pis. 1-26. 

Roberts, H. B. 1953. A new species of decapod crustacean from the Inglis 
member. Bull. Florida Geol. Surv., no. 35, pp. 64-67, pi. 13. 

Ross, A., and R. J. Scolaro. 1964. A new crab from the Eocene of Florida. 
Quart. Jour. Florida Acad. Sci., vol. 27, no. 2, pp. 97-106. 

Department of Geology, University of Florida, Gainesville, 
Florida. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



SURVIVAL POTENTIAL OF PIRANHAS IN FLORIDA 
Martin A. Moe, Jr. 

The presence of piranhas in Florida as aquarium fishes has cre- 
ated some apprehensive speculation on the possibility of these 
characinid fishes becoming established in our native fresh waters. 
The tales of Theodore Roosevelt in his book of 1914, Through the 
Brazilian Wilderness, first popularized the depredations of the pi- 
ranha and established their future reputation. Since that time, 
many writers of both scientific and popular works have eloquently 
described the innate ferocity and feeding frenzies of large schools 
of piranhas. It is the purpose of this paper to compile available 
pertinent information and to attempt rational assessment of the im- 
plications of their importation into Florida. Definitive conclusions 
are not possible because the basic biology and life histories of the 
piranhas are not known, fish ecology in the fresh waters of South 
America has been studied but little, and the taxonomy of piranhas 
is in confusion. Suggestions concerning potential for survival in 
Florida waters can be made but must not be construed as the results 
of extensive biological research. 

There are apparently two major, and rather generalized, divi- 
sions of the piranha genus, Serrasalmus. First of these includes the 
pirambebas, which are usually not considered dangerous and which 
have concave forehead profiles and sharp snouts. The second 
group contains the piranhas, which are considered dangerous to 
man and have slightly decurved or convex foreheads and wide, 
blunt snouts. The major concern of this paper is with the latter 
group although the pirambebas cannot be considered entirely 
harmless. Most authors agree that pirambebas should not be 
trusted too far, and Coates (1933) regards both groups of these 
fishes as "equally bloodthirsty." 

Norman's (1929) work is used as the taxonomic basis for the 
species listed in this paper since it is the most widely accepted 
classification. Gosline (1951) lists all the species of the subfamily 
Serrasalminae that have been described since Norman's paper. 
However, Dr. G. S. Myers (personal communication) of Stanford 
University states "Work recently begun but not finished or pub- 
lished, done here by Dr. A. E. Leviton of the California Academy 
of Sciences, shows clearly that Serrasalmus classification as to spe- 



198 Quarterly Journal of the Florida Academy of Sciences 

cies is in confusion. Norman's revision cannot be accepted as to 
species." The piranha group presently consists of Serrasalmus 
natter eri, the most common and widespread species; S. piraya, the 
largest; S. temetzi; and S. niger. Although, as Dr. Myers points 
out, specific groups may not be distinct, it is generally agreed that 
all four forms are dangerous to man and other animals. Norman 



SOUTH AMERICA - Atlantic drainage 



latitude 




general 
distribution 



Serrasalmus 



is 





S. natterei 



10' 



15 S. piraya 
20' 



25' 


S. 


tern 


etz 


30* 








35* 

40° 


S. niger 

a 


45* 




50* 









Fig. 1. General distribution of the genus Serrasalmus and the piranh; 
throughout South America. 



Moe: Survival Potential of Piranhas 199 

lists 12 additional forms of Serrasalmus that are typified by S. 
rhombeus and are considered to be the less dangerous pirambebas. 
Specimens of S. spilopleura in the Lima Campos reservoir have been 
found by Braga (1956) to be the females of S. rhombeus. 

Figure 1 depicts the approximate range of the genus Serrasal- 
mus and the four species of piranhas recognized by Norman. These 
range lines are based on information presented by Eigenmann 
(1915) and Myers (1949) and represent only rough approximations 
since definitive ranges have not been developed. The genus Ser- 
rasalmus occurs throughout the Atlantic drainage from just south 
of the Maritime Andes of Venezuela to the estuary of the Rio de la 
Plata, with the exception of the short coastal streams between 
southern Uruguay and the Rio San Francisco. The range, shown 
in Figure 1, covers most of the South American continent extend- 
ing from 10° north latitude to 35° south latitude. S. natter eri 
usually occurs throughout this range with the exception of the 
Rio San Francisco which has S. piraya as its only species of piran- 
ha. Eigenmann and Eigenmann (1891) record S. nattereri south- 
ward to the Rio de la Plata. S. ternetzi is restricted to the Rio 
Paraguay and probably the Rio Parana. S. niger has been taken 
only from the area of British Guiana, and is not common in col- 
lections. Various species of pirambebas are found throughout the 
generic range, although S. rhombeus appears to be the most com- 
mon and widespread. Myers always found piranhas and piram- 
bebas in the same stream basins. In a more recent study, Braga 
(1961) found the incidence of pirambebas to be 9.5 times greater 
than piranhas in a study of the Poco da Cruz dam basin. 

Natural History 

Fishes of the genus Serrasalmus are relatively unspecialized 
carnivores, thus are able to adapt to a changing food supply. In 
a study on the feeding habits of the pirambeba, S. rhombeus, Braga 
(1954) states "the pirambeba rarely attacks man or terrestrial ani- 
mals" which implies that attacks have taken place. He found that 
the major portion of their diet consisted of shrimp and fishes, 
although many other animals and small amounts of vegetation 
were included. Eigenmann (1915) has described and pictured the 
dentation of both groups of Serrasalmus and Myers (1949) mentions 
that their jaws are well adapted for neatly removing the precise 



200 Quarterly Journal of the Florida Academy of Sciences 

amount of flesh required for one swallow. Pieces of almost any 
type of flesh, including live fishes, generally constitute their diet 
in an aquarium. It is certain that non-aquatic animals form some 
unknown and variable percentage of their diet, but the primary 
source of nutrition resides in their own environment. Leo W. 
Baumer, tropical fish dealer in Iquitos, Peru, remarks (personal 
communication) that piranhas feed mostly on other fishes, start- 
ing at the tail and working toward the head, and occasionally 
engage in cannibalism. 

There are three reports of Serrasalmus spawning in captivity 
in the United States. Braker (1960) first reported the spawning 
of S. spilopleura in a 1200 gallon tank at the John G. Shedd Aqua- 
rium. The spawning took place in water of 7.6 pH and 78 °F. 
(approximately 24° C). The fish were seven inches long and had 
been in captivity three years before they spawned. Mr. Kyle 
Swegles, owner of the Rainbow Aquarium in Chicago, is reported 
by the Tropical Fish Hobbyist (Anonymous, 1963) to have spawned 
S. natter eri. Mr. Swegles stated that his piranhas required a diet 
of five mice to be kept in prime spawning condition. The fish 
spawned once every two weeks, released up to 5,000 adhesive 
eggs, and showed careful parental guarding of the eggs. S. niger 
was also spawned at the John G. Shedd Aquarium by Braker 
(1963). In both instances reported by Braker, the males chased 
away the females after spawning and diligently guarded the eggs. 
The S. niger that spawned were about three years of age and 10 
to 12 inches long. Young S. spilopleura were reported by Braker 
(1963) to reach a length of 1% inches in 3V2 months under aquarium 
conditions. Braker (personal communication) reports that two 
specimens of S. niger at the Shedd Aquarium lived more than 20 
years and exceeded 18 inches and 5V2 pounds when they died. 
Braga (1956) states that the pirambeba, S. rhombeus, is a fish of 
"intense sexual activity" and Leo W. Baumer (op. cit.) remarks 
that small piranhas are abundantly collected during November, 
December, and January near Iquitos, Peru. Identifications of these 
fishes, particularly S. niger, are questionable because of the taxo- 
nomic confusion in the genus Serrasalmus. Longevity, reproduc- 
tive cycles and growth patterns are not known in the wild. 

Piranhas are found in most aquatic environments within their 
geographic range in South America. Myers states that they stay 
near the bottom of the quieter, deeper waters; and Leo W. Baumer 



Moe: Survival Potential of Piranhas 201 

remarked to me that piranhas seem to prefer the clean waters 
of small, slow-moving streams to the murky waters of the Amazon. 

Danger to Man 

Extensive, well-documented accounts of piranhas attacking man 
and other terrestrial and aerial animals can be found in several 
of the references cited in this paper and it would serve no purpose 
to list them here. Myers (1949) mentions that an attraction to un- 
usual water disturbances and a tendancy to nibble on larger ani- 
mals are behavioral traits of many characins. The primary dan- 
ger to larger animals stems from the fearless attack of half-grown 
and adult piranhas in groups often approaching hundreds of in- 
dividuals. One instance of the destruction of a steer by piranhas 
has been photographically recorded by Cognac (1963) in the Bra- 
zilian state of Matto Grosso. 

Infrequent but dramatic instances have done much to build 
the notorious reputation of these fishes, and undoubtedly the tales 
gain many embellishments as they are retold. Schultz (1960) states 
that in 20 years of collecting tropical fishes in South America he 
has come in contact with only seven persons that have been bitten 
by piranhas; and Leo W. Baumer (personal communication) also 
states that in his years of collecting tropical fishes he has not heard 
a first-hand account of an attack on man. I. B. A. Rokes, manager 
of South American Fish Exporters in Iquitos, Peru, has encount- 
ered only two instances of piranha bites in 18 years of collecting 
tropical fishes (personal communication). He also believes, based 
on his experience, that the danger to man from the Amazon pi- 
ranhas is exaggerated. Daniel M. Troth, Biological Aide with 
the Florida Board of Conservation, dove among piranhas in the 
waters of Surinam and reports that the individuals and small 
groups he encountered seemed timid and avoided him. The na- 
tives indicated to him that the waters were only dangerous at 
certain times. Mr. Troth's comments agree with observations on 
individual specimens kept in aquaria by Mr. Ross Socolof, Gulf 
Fish Hatchery, Palmetto, Florida. Fontenele (1963) presents pho- 
tographs of the results of two bites by piranhas; and The Aquarium 
(Anonymous, 1935) reports on three bites suffered by humans from 
captive piranhas, most likely S. natter eri. Present knowledge of 
these fishes is too incomplete to allow behavioral analysis; but the 



202 Quarterly Journal of the Florida Academy of Sciences 

general statement can be made that the piranhas, and perhaps the 
pirambebas, are potentially dangerous (certainly under conditions 
of capture); and at certain times in localized areas they are very 
dangerous. 

Potentials for Acclimatization 

According to Briggs (1958) "the freshwater fish fauna of Florida 
owes its relationship and origin to the fauna of the continental 
United States." This native fauna is not rich, since Florida has 
recently emerged on the geologic scene, and most temperate spe- 
cies have had limited success in moving and adapting to a low 
land with a subtropical climate. The Florida peninsula extends 
from about 25° to 30° north latitude. Its southerly location, com- 
bined with the effects of the warming Florida Current along the 
lower east coast, produces a tropical climate in the southern tip of 
Florida and a subtropical climate through central Florida. Sim- 
ilarities of climate suggest that piranhas from south-central South 
America may be able to adapt to Florida's climate. 

Low water-temperature tolerance appears to be the critical 
factor in the possible establishment of piranhas in Florida. The 
importance of pH can be minimized since Schultz (1960) reports 
that piranhas occur naturally in acid to slightly alkaline waters 
and Braker (1960) spawned S. spilopleura in pH of 7.6. Black 
and Brown (1951) state that Florida's deep ground waters are 
usually alkaline, varying between 7.0 and 8.0 in pH. Surface 
waters fluctuate rapidly in pH value because of the actions of 
living organisms. Some surface waters almost always remain 
between 6.0 and 7.0 in pH due to the presence of free carbon 
dioxide and organic acids from runoff. The author tested 11 loca- 
tions in various canals of south Florida between Homestead and 
Fisheating Creek on U. S. Highway 27 for pH and temperature 
on March 8, 1964. The pH ranged from 6.1 to 7.3 with the ma- 
jority falling between 7.1 and 7.2. Surface temperatures ranged 
from 24°C. to 28°C. and averaged about 25°C. 

The surface water temperature in the vicinity of Fort Lauder- 
dale, Florida during 1960 and 1961 has been discussed by Clugs- 
ton (1964). December 1960 and January 1961 exhibited the cool- 
est readings during this period. The temperature dropped to 
60°F. (15°C.) for several days during these months, although 
the average minimum and maximum temperatures were 65 °F. 



Moe: Survival Potential of Piranhas 203 

(18 C C.) and 68°F. (20°C). During the summer months the tem- 
perature usually ranged between 80°F. (26° C.) and 86°F. (30°C). 
Black and Brown (1951) state "The temperature of the surface 
waters, of course, varies with the season and reflects in general 
the temperature of the surroundings." Table 1 applies this cri- 
terion to obtain a general comparison of the southern range of 
the piranha and southern Florida. The temperatures listed are 
the long term mean average for the coldest and warmest months 
of the year. Paraguay is bisected at 25° south latitude and the 
temperatures are similar to those of south Florida. (The piranhas 
of Paraguay are generally considered quite dangerous). These 
data suggest that, in respect to temperature, piranhas derived 
from the southern portion of their geographic range would be 
able to tolerate the temperature range of southern Florida. 

TABLE 1 
Mean average temperature for coldest and warmest months of year 

Annual 
Location Summer Winter Average 

Miami Airport, Florida 82.9°F 68.7°F 75.7°F 

Belle Glade Exp. Sta., Florida 80.6°F 63.8°F 72.4°F 

Rio de Janeiro, S. A. 68.7°F 74°F 

Paraguay, S. A. 81 °F 64°F 74°F 

Uruguay, S. A. 71 °F 50°F 

Buenos Aires, S. A. 73.6°F 49°F 61 °F 



Florida data from U. S. Dept. of Commerce, Weather Bureau, Climatolog- 
ical Data, Florida for 1958, vol. 57, no 13, pp. 196-205. 

South American data from Encyclopaedia Britannica, 1953, vol. 4, pp. 
338-339; vol. 17, p. 258; vol. 19, pp. 314-315; vol. 22, pp. 904-906. 

Man has altered and controlled the discharge of fresh water in 
southern Florida by creating an extensive interconnecting net- 
work of canals, many of which are deep and filled with clean, 
slow-moving water. Such deeper water does not change in tem- 
perature and oxygen content as rapidly as do shallow waters; 
thus seasonal changes are more gradual and daily variations less 
extreme. 



204 Quarterly Journal of the Florida Academy of Sciences 

The tolerances of piranha to temperature, rate of temperature 
change, and salinity are unknown in their native waters. Mr. 
Clarence Thompson, owner of the Aquarium at Key West, Florida, 
verbally reports holding a piranha for five years in unfiltered tap 
water at about 70 °F. (approximately 21 °C). Reports from the 
public on this and other piranhas in aquaria first called our atten- 
tion to the presence of these fish in Florida. It appears that the 
possibility of accidental or intentional introduction is limited 
to fresh waters. 

Animals that inhabit a wide range of biotopes and have a gen- 
erally unspecialized diet would be most likely to survive in a simi- 
lar but alien environment. Piranhas, particularly S. nattered, 
fulfill these requirements. Parental care of the eggs and prolifi- 
cacy further increase the survival potential. 

Southern Florida's fresh waters already support a number of 
euryhaline and freshwater carnivores such as snook, tarpon, bass, 
and gar. Their prey, supplemented by various invertebrates as 
well as the carnivores themselves, would form a ready food sup- 
ply for the aggressive piranha. 

Freshwater fishes from the tropical regions of the world are 
not new to Florida's waters. At least five species of fishes im- 
ported for the aquarium trade are now living and reproducing in 
our native waterways. The cichlid, Tilapia heudeloti, has become 
established in the Tampa area and is reported by Springer and 
Finucane (1963) to have entered the commercial fish catch. Its 
native range is western Africa, roughly from 15° north latitude 
to 10° south latitude, and Sterba (1962) reports its temperature 
range as "22 to 24 °C, resistant to temperatures not far below 
20 °C." T. heudeloti has survived temperatures much lower than 
20° C. in Florida waters. Socolof (1963) reports the establishment 
of Plecostomus sp., also in the Tampa area. This fish has the same 
general range in South America as the southern groups of Serrasal- 
mus. The establishment of two tropical cichlids and one cyprino- 
dont in the waters of southeastern Florida is reported by Dr. C. 
Richard Robins of the Institute of Marine Science, University of 
Miami (personal communication). The cichlids are Astronatus 
ocellatus, which ranges from Brazil to northern Argentina, and 
Hemichromis sp., widespread in central Africa. The cyprino- 
dont, Belonesox belizanus, is from Central America between 9° to 
21° north latitude. An animal in temperate latitudes must be 



Moe: Survival Potential of Piranhas 205 

able to survive the severest of winters in order to continue exist- 
ence in that climate. In 1963 these fishes survived one of the cold- 
est winters ever recorded in Florida. 

Ten thousand juvenile Cichla ocellaris were released recently 
by the Florida Game and Fresh Water Fish Commission in the 
Fort Lauderdale area in an effort to investigate the results of 
establishing this South American sport fish. There is no record 
of any established characin in Florida, although the range of the 
family extends northward into Texas. The possible effects of 
these newly introduced species on the existing community structure 
is not known; however, their successful establishment proves the 
ability of some extrinsic tropical species to adapt to the Florida 
environment. 

Importation 

Herald (1961) mentions that S. nattered, the commonest of the 
piranhas, is the species most often imported and exhibited in the 
United States. The pirambebas are also often exhibited under the 
common name piranha. S. rhombeus is probably the pirambeba 
most often imported, as it is the most widespread species. 

Piranhas entered the aquarium trade primarily as novelties, and 
are frequently displayed by dealers and exhibitions to attract the 
public. Ross Socolof informs me that the greatest trade in pi- 
ranhas occurred during 1960 and 1961 and that the demand is 
very limited at the present time. In comparison with other fishes, 
very few piranhas are exported by Leo W. Baumer of Iquitos, 
Peru (op. cit). Late in the summer of 1962 the Florida Game 
and Fresh Water Fish Commission requested that the U. S. Cus- 
toms service aid in enforcing the statute restricting the importation 
of piranhas. Only holders of permits issued by the Commission 
are allowed to handle or display piranhas. According to A. D. 
Aldrich (personal communication) Director, Game and Fresh Wa- 
ter Fish Commission, permits had been issued to dealers for the 
transshipment of 2,738 piranhas and 30 possession permits had also 
been issued to legitimate public exhibitions as of February 4, 1964. 
Herald (1961) mentions that before the first spawning of piranhas 
in captivity no laws against import were considered, since it was 
assumed that permanent populations could not be established. 

The shipment of live piranhas of large juvenile or adult size 
presents difficulties that have been well described by Atz (1955). 



206 Quarterly Journal of the Florida Academy of Sciences 

Problems of collection, shipment (they must be tranquilized and 
packed individually), and handling all raise the price and low- 
er the demand. Almost all shipments of tropical fish are trans- 
ported by commercial airlines. Numerous fish are placed in large 
plastic bags that usually hold several quarts of water, often a 
tranquilizer, and an oxygen source. This bag is held in a heavy 
styrofoam or cardboard box during shipment. Additional tranquil- 
izer and oxygen may be added on long trips. Most of the piranhas 
that are imported are young juveniles that find their way into the 
shipments of other characins. Young piranhas are so similar in 
shape and color to many of their vegetarian relatives that only an 
experienced ichthyologist can distinguish them. They often are 
not recognized until they reach the tanks or outdoor ponds of the 
wholesaler. Miami and Tampa are the only Florida ports of entry 
for tropical fishes, and the major fish farms are located in these 
areas. Earl S. Herald, Superintendent-Curator of Steinhart Aquar- 
ium, in a letter to Daniel H. Janzen of the Bureau of Sport Fish- 
eries and Wildlife, stated that piranhas are still readily available 
on the tropical fish market under a variety of Spanish, Portuguese, 
and native names. For these reasons importation of piranhas is 
difficult to regulate. 

Discussion 

Man has always endeavored to change and control nature. He 
often has introduced extrinsic life forms in order to produce more 
food, preserve old customs, or obtain unusual ornamentation. On 
occasion he has introduced unknowingly exotic plants and animals 
through the mere fact of his presence. These introductions fre- 
quently have aided man's efforts to better his environment but in 
many cases their effect was detrimental to both the local biota 
and his labors. 

Most of the fishes imported for the tropical fish trade originate 
from the central and northern portions of South America. These 
fishes are adapted to equatorial conditions even if the species ex- 
tends into temperate South America. Whether or not these fish 
all possess the potential to survive in a more temperate climate 
is a moot point. Under favorable conditions, a mild winter, ac- 
cidentally liberated piranhas may be able to survive and repro- 
duce, and then natural selection would favor those individuals 
with the lowest temperature tolerances. Even if these fishes were 



Moe: Survival Potential of Piranhas 207 

able to withstand a few degrees of salinity, the problems devel- 
oped by their importation are restricted to Florida's native fresh 
waters. 

The sole beneficial aspect of the presence of piranhas is that 
they are easy to catch with the proper equipment and are of good 
flavor with few bones. 

These fishes are not only a danger to men and livestock, but 
they also create havoc with fishermen by destroying hooked fish 
and fish nets. Their undesirability is evidenced by the efforts of 
the Brazilian government to eradicate them from the important 
inland water areas. Fontenele (1963) describes these efforts and 
their success. Many thousands of dollars have been spent in poi- 
soning these fishes with rotenone and in building dams to prevent 
reinfestation. 

On the basis of the information presented above, I believe that 
accidental establishment under the present conditions of import 
and control is possible, but unlikely. On the other hand, if piran- 
has were desired in southern Florida, I would predict that their 
establishment could be successfully achieved. 

The genus Serasalmus would not be a worthwhile addition to 
the fresh waters of Florida because of their danger to man, and 
every effort should be made to prevent introduction. The vast in- 
ter-connected waterways of south Florida would make eradica- 
tion costly and success improbable. 

Addendum 

Recent correspondence from R. Adhemar Braga, biologist with 
the Dept. Nacional de Obras contra as Serras, Divisao de Pesca 
e Piscicultura, of Brazil, supplements the information presented 
above. 

Piranhas are commonly found from sea level to heights of 600 
meters (1,968 feet) and extend southward in Brazil to about 27° 
south latitude in the tributaries of the Paraguay River. They are 
always considered a nuisance and are being eradicated from many 
waterways in northeast Brazil. The economic value of the piranha 
is very low in Brazil. 

Data extracted from Menezes (1960) show maximum and min- 
imum monthly water temperatures from June 1941 through July 
1943 for the Lima Campos dam on the Parnahiba River, approxi- 



208 Quarterly Journal of the Florida Academy of Sciences 

mately 7° south latitude. The minimum temperature of 20.0° C 
(approximately 68 °F) occurred in August 1942, and the maximum 
for that month was 27.8 °C (approximately 81 °F). The maximum 
temperature (32.2 °C, approximately 95 °F) occurred in March 1943, 
and the minimum for that month was 27.0 °C or about 80 °F. 
These temperature extremes, even though taken much closer to the 
equator, do not differ greatly from those reported by Clugston 
(1964) for south Florida. Piranhas spawn in this area during all 
months of the year, apparently with no special ecological require- 
ments. 

Acknowledgments 

I am indebted to those individuals who gave me of their time 
and knowledge through correspondence on various facets of this 
paper. They are acknowledged in the text. My thanks go also 
to Doctors James W. Atz, George K. Reid, Victor G. Springer, 
and Stanley H. Weitzman, who critically reviewed the manuscript 
and offered many meaningful revisions. Mr. Robert M. Ingle, Di- 
rector of Research, Salt Water Fisheries Division, Florida Board 
of Conservation, suggested this line of inquiry and provided coun- 
sel and encouragement. 

Literature Cited 

Anonymous. 1935. New member initiated in the piranha club. Aquarium, 
vol. 4, no. 7, p. 146. 

. 1963. Piranhas spawned by famous Chicago hobbyist-dealer. Trop- 
ical Fish Hobbyist, vol. 11, no. 9, pp. 75-76. 

Atz, James W. 1955. Rare black piranha comes to New York. Aquarium 
Journal, vol. 26, no. 1, pp. 3-7. 

Black, A. P., and Eugene Brown. 1951. Chemical character of Florida's 
waters 1951. Florida Div. Water Survey & Research, Paper no. 6, 
119 pp. 

Braga, R. Adhemar. 1954. Alimentacao de pirambeba, "Serrasalmus rhom- 
beus" (L., 1766) Lacepede, 1803, no acude Lima Campos, Ico, Ceara 
(Ostariophisi, Characidae, Serrasalminae). Rev. Brasil. Biol., vol. 14, 
no. 4, pp. 477-492. 

. 1956. Carater sexual secundario em pirambeba "Serrasalmus rhom- 

beus" (L., 1766) Lacepede, 1803. Rev. Brasil. Biol, vol. 16, no. 2, 
pp. 167-180. 



Moe: Survival Potential of Piranhas 209 

. 1961. Erradicao de piranhas no acude publica "Poco oa Cruz" 

(Inaja, Pernambuco) 1. Reconhecimento oa bacia hidrografica (1). 
Bol. Museu Nacional, Zoologia, no. 226, 32 pp. 

Braker, William P. 1960. Bill Braker's tank talk. Aquarium, vol. 29, no. 
1, pp. 6-10. 

— . 1963. Bill Braker's tank talk. Aquarium, vol. 32, no. 10, pp. 12-14. 



Briggs, John C. 1958. A list of Florida fishes and their distribution. Bull. 
Florida State Mus., vol. 2, no. 8, pp. 223-318. 

Clugston, James P. 1964. Growth of the Florida largemouth bass, Mi- 
cropterus salmoides floridanus (LeSuer), and the northern largemouth 
bass, M. s. salmoides (Lacepede), in subtropical Florida. Trans. Amer. 
Fish. Soc, vol. 93, no. 2, pp. 146-154. 

Coates, C. W. 1933. Tropical fishes as pets. Liveright, New York, p. 226. 

Cognac, Marcel. 1963. Piranha. Sports Afield, vol. 150, no. 5, pp. 46-47. 

Eigenmann, Carl H. 1915. The Serrasalminae and Mylinae. Annals Car- 
negie Mus., vol. 9, nos. 3-4, pp. 226-272. 

Eigenmann, Carl H., and Rosa S. Eigenmann. 1891. A catalogue of the 
fresh water fishes of South America. Proc. U. S. Nat. Mus., vol. 14, 
pp. 1-81. 

Fontenele, Osmar. 1963. Eradication of piranha in inland waters. Comm. 
Fish Review, vol. 25, no. 3, pp. 46-50. 

Gosline, W. A. 1951. Notes on the characid fishes or the subfamily Serras- 
alminae. Proc. Calif. Acad. Sci., vol. 27, no. 2, pp. 17-64. 

Herald, Earl S. 1961. Living fishes of the world. Doubleday & Company, 
New York, 304 pp. 

Menezes, R. S. 1960. Contribuicao para o estudo da pesca no acude "Lima 
Campos." Bol. Dept. Nac. Obras contra Seras, vol. 22, no. 9, pp. 
1-110. 

Myers, George S. 1949. A monograph on the piranha. Aquarium, vol. 20, 
no. 2, pp. 52-61; no. 3, pp. 76-85. 

Norman, J. R. 1929. The South American characid fishes of the genus Ser- 
rasalmus Lacepede. Proc. Zool. Soc. London for 1928, pp. 781-829. 

Schultz, Harald. 1960. Piranhas, fact and fiction. Tropical Fish Hobby- 
ist, vol. 9, no. 1, pp. 33-59. 

Socolof, Ross. 1963. Iquitos: fish collecting capital of the world. Tropical 
Fish Hobbyist, vol. 11, no. 9, pp. 74-78. 



210 Quarterly Journal of the Florida Academy of Sciences 

Springer, Victor G., and John H. Finucane. 1963. The African cichlid 
Tilapia heudeloti Dumeril, in the commercial fish catch of Florida. 
Trans. Amer. Fish Soc, vol. 92, no. 3, pp. 317-318. 

Sterba, Gunther. 1962. Freshwater fishes of the world. Viking Press, 
New York, 878 pp. 

Florida Board of Conservation Marine Laboratory, St. Peters- 
burg, Florida. Contribution No. 79. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



NEW SUBSPECIES OF LEIOCEPHALUS FROM CUBA 
Albert Schwartz 

The taxonomic status of specimens of Leiocephalus from east- 
ern Cuba is in question. Some of these lizards were tentatively 
regarded by me as L. cubensis; later I regarded them as of the 
species L. stictigaster but did not assign them to subspecies 
(Schwartz, 1959, 1960). I had hoped to secure fresh comparative 
material during the summer of 1960, when I collected in Cuba 
under National Science Foundation Grant G-6252. This expecta- 
tion was at least partly successful, since now the population of 
L. stictigaster from the serpentine savannas in Camagiiey Province 
is well known. No additional material has been taken in Oriente, 
however. A day's trip to Cayo Sabinal, off the north coast of Cam- 
agiiey, likewise produced a series of L. stictigaster which repre- 
sents a new insular form. 

I wish to acknowledge the assistance of the National Science 
Foundation in subsidizing my collecting in Cuba. During the 
summer of 1960 I had the enthusiastic and capable assistance of 
Messrs. Ronald F. Klinikowski, David C. Leber, and James D. 
Smallwood; Messrs. Klinikowski and Leber had National Science 
Foundation Undergraduate Research Participation grants at that 
time. Mr. Klinikowski has executed the figures for the present 
paper as well. For comparative material I wish to thank Dr. Ernest 
E. Williams, Museum of Comparative Zoology (MCZ), who has 
allowed me to reexamine some pertinent old Oriente material, and 
Sr. Miguel Jaume, of the Museo y Biblioteca de la Zoologia de la 
Habana (MBZH) for like permission. Our visit to Cayo Sabinal 
was made possible through the cooperation of Sr. Carlos Betan- 
court. 

L. stictigaster occurs in two general areas: in Pinar del Rio 
Province and the Isla de Pinos, and in Camagiiey and Oriente 
provinces. There are two subspecies known from Pinar del Rio 
(stictigaster and sierrae), two from the Isla de Pinos (exotheotus 
and astictus), and another from a very restricted area in Camagiiey 
(lucianus). L. cubensis, the larger species to which stictigaster is 
related and with which it was long confused, is not known to be 
sympatric with stictigaster at any one locality. The subspecies 
lucianus, for example, whose range is included within the presumed 



212 Quarterly Journal of the Florida Academy of Sciences 

range of L. c. cubensis, does not occur with cubensis, although it 
does occur with or immediately adjacent to the species L. carinatus 
and macropus. Of the two new subspecies described herein, nei- 
ther is known to occupy territory with any other Leiocephalus; 
on Cayo Sabinal we encountered only stictigaster and in the serpen- 
tine savannas, no other Leiocephalus was seen or taken except this 
same species. 

Leiocephalus stictigaster parasphex, new subspecies 

Type. American Museum of Natural History (AMNH) 92153, 
an adult male, from Playa Bonita, east end Cayo Sabinal, Cam- 
agiiey Province, Cuba, taken 31 July 1960, one of a series collected 
by Ronald F. Klinikowski, David C. Leber, and James D. Small- 
wood. Original number 9596. 

Paratypes. AMNH 92154-73, same data as type. 

Diagnosis. A subspecies of Leiocephalus stictigaster character- 
ized by longitudinally lined dorsum in which the dorsal fields are 
solid brown, lateral fields brown with red or orange flecking in 
males, throat and venter pale yellow, venter without dark spots 
but with red or orange spots laterally, throat clouded grey with 
a linear pattern present but not distinct, and parietal scales usually 
not in contact. 

Distribution. Known only from the type locality but presumed 
to occur throughout Cayo Sabinal. 

Description of type. An adult male with the following measure- 
ments (in mm) and counts: snout-vent length 65, tail 67 (distal 
two-thirds regenerated), snout to anterior border of tympanic open- 
ing 15.3, head width 12.2, supraocular scales 6/6, loreals 5, tempor- 
als 10, enlarged auricular scales 5/4, median head scales 4, pre- 
frontal row complete 3 scales, frontoparietal row destroyed, semi- 
circles unknown, parietal contact unknown, dorsal crest scales occi- 
put to vent 50, dorsal crest scales occiput to axilla 16, scales around 
one half body at mid-body 24, fourth toe subdigital tricarinate 
scales 23/23. 

Coloration of type. The dorsal coloration of the type is gen- 
erally dark brown with orange sides, the pigments arranged in the 
typical zoned pattern of stictigaster as follows: Zone 1 is narrow 
and brown and restricted to the dorsal crest scales. Laterally, 
Zone 2 is dull tan and rather sharply set off from Zone 3, the dorso- 



Schwartz: New Lizards from Cuba 213 

lateral fields, which are very dark rich brown. The dorsolateral 
fields are again sharply differentiated from Zone 4 which is wide 
and tan, and extends from the temporal region posteriorly onto 
the unregenerated proximal third of the tail. Zone 5 is brown 
with much vivid red and orange flecking; it begins posterior to the 
eye and continues to the groin. There are no longitudinal dark 
brown dashes in Zone 5. Zone 6 is cream, very bold and distinct, 
beginning below the eye, continuing over the tympanic opening 
and reaching the groin posteriorly. Below Zone 6 is a bright 
orange longitudinal region (Zone 7), which is reddish-orange and 
very brilliantly differentiated from both Zone 6 above and the 
pale yellow ventral coloration below. The dorsal surface of the 
head is solid brown without darker brown or black pigmentation. 
The postorbital blotch is absent, the temporal region having the 
same brown coloration as Zone 5. The dorsal surfaces of the 
limbs are brown with some orange and tan flecking: neither fore- 
nor hind limbs have any indication of dark flecks or dashes. The 
unregenerated portion of the tail lacks chevrons or other dark 
dorsal markings but does show the caudal extension of tan Zone 4. 
The throat is diffusely marked with dark gray on a clouded grayish- 
yellow ground color; there are two large V's, their apices directed 
forward, and a posterior pair of paramedian lines, superimposed 
on almost a gray reticulum, which in turn is only moderately dis- 
tinct from the clouded grayish-yellow ground color (Fig. 1). The 
gray reticulum extends posteriorly onto the chest. The remainder 
of the belly is immaculate pale yellow, as is also the underside of 
the limbs and tail. 

Variation. In snout- vent length, ten males (type and para types) 
average 63.7 (57-75), and eleven paratypic females average 55.1 
(50-59). Dorsal crest scales in occiput- vent length (combined data 
for both sexes) average 52.1 (46-58), and dorsal crest scales in occi- 
put-axilla length average 19.5 (16-22). One-half scales at mid-body 
average 22.9 (21-27), loreals 5.2 (3-10), temporals 11.4 (10-14), fourth 
toe subdigital tricarinate scales 23.6 (22-26). The parietals are 
more often not in contact (78 per cent), and the semicircles are more 
often in contact (80 per cent). The median head scales vary from 
4 to 7 (mode 4). The prefrontal row is complete in all examples, 
and varies from 3 to 6 scales (mode 3). The frontoparietal row is 
usually complete (twelve of fifteen lizards) and has 4 to 9 scales 
(mode 5). 



214 Quarterly Journal of the Florida Academy of Sciences 

The para types include nine males, all adult. In dorsal colora- 
tion and pattern these agree very closely with the type, all having 
the distinct and bold longitudinal zones, the dorsolateral fields dark 
brown, the pale tan or cream lines, and the brightly colored lower 
sides (Zone 7). Some adult males have the dorsolateral fields 
brown, heavily specked with red or orange, and these colors always 
form a prominent part of the lateral fields (Zone 5) and the lower 
accessory Zone 7. The throat pattern in males is variable in both 
extent and intensity. The ground color is always clouded with 
grey. The two gray V's may be entire or much fragmented, and 
the two paramedian posterior lines are likewise variable. These 
major pattern elements may be superimposed upon a gray to dark 
gray reticulum, which may as well combine with the V's and para- 
median lines, to give a uniform dark gray reticulum. The gray 
throat pattern may extend rather far posteriorly onto the chest or 
even down onto the sides of the abdomen; in the latter case the 
dark gray spots are replaced by isolated red or orange scales, which 
follow the same general trend as the dark gray, more anterior scales 
and are a continuation of the gray pattern posteriorly. The bellies 
of all specimens are immaculate. 






Fig. 1. Leiocephalus stictigaster parasphex, new subspecies, ventral view 
of throat; type, AMNH 92153. Fig. 2. Leiocephalus stictigaster lucianus, 
ventral view of throat; AMNH 92174, 1 mi. E Playa Santa Lucia, Camaguey 
Prov., Cuba. Fig. 3. Leiocephalus stictigaster ophiplacodes, new subspecies, 
ventral view of throat; type, AMNH 92771. 



The females resemble the males dorsally, except that there is 
a distinct reduction of amount of red and orange in all fields, these 
colors being completely absent except in large females. The ven- 
tral surface is pale yellow in all; there may be dull orange spots 



Schwartz: New Lizards from Cuba ,-::::.. 215 

on the venter, or it may be immaculate. The throat pattern re- 
sembles that of males except that it is much more diffuse. The 
two V's may disappear almost completely, leaving only dark frag- 
ments, or they may be joined randomly to one another by dark 
reticular markings. The two paramedian lines retain their integ- 
rity to a greater extent and may even be paired laterally with an- 
other pair of dark longitudinal lines which are also occasionally 
indicated in males. The throat ground color is always a dirty 
yellow. Only one female shows any indications of dark leg dots 
or markings on the hindlimbs, and none, like the males, has any 
indication of dark dashes in the lateral fields. 

Comparisons. The range of L. s. parasphex is removed from 
that of L. s. lucianus by a narrow channel with a maximum depth 
of fifteen fathoms. Consequently, parasphex needs comparison 
principally with lucianus. The two subspecies are very different in 
coloration and pattern. The throat pattern differences are at once 
obvious. Both sexes of lucianus have bold black throat markings 
(Fig. 2) that are almost always complete (two V's and a pair of para- 
median lines as in parasphex) and, even when reduced in extent, 
are not reduced in intensity. In lucianus there is no indication of 
throat clouding nor a gray throat reticulum. The dorsal pattern 
of lucianus lacks red or orange as a chromatic component, the in- 
sular race having much more brightly colored back and sides. 
All male lucianus have brown ventral dots which extend onto the 
underside of the hind limbs and along the sides of the tail. These 
dots are absent in parasphex. The throat pattern of lucianus often 
includes a short transverse median bar anterior to the apex of the 
throat V; this item is absent or at best very obscure in parasphex. 
The females of lucianus, besides having a much more clear-cut 
throat pattern than those of parasphex, likewise have the venter 
more heavily and uniformly dotted with brownish. 

L. s. parasphex and lucianus are close in all scale counts. There 
are no striking differences in number of dorsal crest scales, either 
between the occiput and vent or occiput and axilla, although 
parasphex averages very slightly higher (52.1 vs. 51.6) in occiput- 
vent scales and somewhat lower (19.5 vs. 22.0) in occiput-axilla 
scales. Both have the semicircles more often complete than incom- 
plete. The major scale difference is in parietal contact; lucianus 
usually (73 per cent) has the parietals in contact whereas parasphex 
usually (78 per cent) has them not in contact. 



216 Quarterly Journal of the Florida Academy of Sciences 

Remarks. The northern coast of Camagiiey Province has, in 
addition to many small cays and islets, a series of large cays, par- 
alleling the coast for about 190 kilometers (Fig. 4). These cays 
include, from west to east, Cayos Coco, Romano, Guajaba, and 
Sabinal. Sabinal is thus the easternmost of the series and is closest 
to the mainland, being separated only by two very narrow chan- 
nels and the Bahia de Nuevitas. Of the other cays, Romano is 
tied to the mainland by a narrow isthmus. The herpetological 
fauna of these cays is completely unknown except for that of Sabi- 
nal. Whether L. stictigaster occurs on the other three cays in this 
chain is unknown. 

Etymology. From Greek para (along side of) and sphex (wasp), 
in reference to the abundant wasp population that makes collect- 
ing somewhat hazardous on Cayo Sabinal. 




Fig. 4. Eastern Cuba, including the provinces of Camaguey and Oriente, 
showing known distribution of subspecies of L. stictigaster, as follows: 1) para- 
sphex; 2) lucianus; 3) ophiplacodes; 4) Bayamo specimens. 



Schwartz: New Lizards from Cuba 217 

Leiocephalus stictigaster ophiplacodes, new subspecies 

Type. AMNH 92771, an adult male, from 2.7 mi. SE Bonao, 
Camagiiey Province, Cuba, taken 3 August 1960, one of a series 
collected by Ronald F. Klinikowski and James D. Smallwood. 
Original number 9666. 

Paratypes. AMNH 92770, 92772-78, same data as type; AMNH 
92779-80, 0.1 mi. SE Bonao, 3 August 1960, A. Schwartz, J. D. 
Smallwood; AMNH 92781-82, 1.5 mi. SE Bonao, 3 August 1960, 
R. F. Klinikowski, J. D. Smallwood; MCZ 59229, ca. 20 km N 
Camagiiey, Camagiiey Province, Cuba, 21 August 1959, R. Mo- 
lina, E. E. Williams, R. Ruibal; MCZ 59335, south of west end of 
Sierra de Cubitas, Camagiiey Province, Cuba, 1959, R. Molina, 
E. E. Williams, R. Ruibal. 

Diagnosis. A subspecies of Leiocephalus stictigaster character- 
ized by longitudinally lined dorsum in which the dorsal fields are 
solid brown, orange spots on sides of abdomen large and prom- 
inent and blending into orange lower sides, distinct cream longi- 
tudinal stripes on back and sides, vivid orange groin, venter pale 
yellow with orange to brown dots, cream colored throat with bold 
and distinct black pattern, large size, high number of dorsal crest 
scales in occiput-vent and occiput-axilla distances, and high num- 
ber of midbody scales. 

Description of type. An adult male with the following measure- 
ments and counts: snout- vent length 70, tail 59 with regenerated 
tip, snout to anterior border of tympanic opening 16.5, head width 
12.5, supraocular scales 6/6, loreals 5, temporals 15, enlarged auric- 
ular scales 4/2, median head scales 4, prefrontal row complete 3 
scales, frontoparietal row complete 5 scales, semicircles incomplete, 
parietal contact present, dorsal crest scales occiput to vent 58, dor- 
sal crest scales occiput to axilla 20, scales around one-half body 
at mid-body 28, fourth toe subdigital tricarinate scales 18/21. 

Coloration of type. The dorsal coloration of the type is gen- 
erally dark brown with orange on the side and lower sides, the 
pigments arranged in the typical zoned pattern of stictigaster as 
follows: Zone 1 inconspicuous and blending into Zone 2 which 
is pale brownish and not especially sharply set off from dark brown 
Zone 3, the dorsolateral fields, which in life were rather heavily 
flecked with orange. The dorsolateral fields are fairly sharply 
differentiated from Zone 4, which is cream and extends from the 



218 Quarterly Journal of the Florida Academy of Sciences 

temporal region onto the base of the tail. Zone 5 is brown with 
heavy orange flecking, most pronounced in the groin, which is 
vivid orange in life; Zone 5 lacks conspicuous dark brown dashes. 
Zone 6 is cream and very distinct, beginning below the eye and 
extending to the groin. Below Zone 6 is a bright orange longitud- 
inal region (Zone 7), which was vividly orange in life. The dorsal 
surface of the head is brown without dark brown or black spotting 
or markings. The dorsal surface of the forelimbs is brown, flecked 
with some orange, and the hind limbs were orange dorsally with 
cream spots. The unregenerated portion of the tail has a dorsal 
pattern of about eleven chevrons, with their apices pointed pos- 
teriorly, which follow four pairs of brown spots on the base of the 
tail. The venter is pale cream with a bold black throat pattern, 
consisting of the incomplete remnants of two V's, their apices 
pointed forward, and two pairs of longitudinal lines on the throat 
itself, the paramedian pair much fragmented and barely discern- 
ible, the lateral pair more or less attached to the posterior V 
(Fig. 3), the whole throat pattern blending into some orange spot- 
ting on the chest, and on the sides of the abdomen, where the scat- 
tered orange dots merge with the orange pigment of Zone 7. The 
underside of the limbs and tail is immaculate cream, without any 
indication of dark dots or longitudinal dashes. 

Variation. In snout- vent length, six males (type and paratypes) 
average 67.7 (57-73), and seven paratypic females average 63.3 (58- 
70). Two paratypes are juveniles with snout-vent lengths of 27 
and 33. Dorsal crest scales in occiput-vent length (combined data 
for all specimens) average 57.2 (51-60), and dorsal crest scales in 
occiput- axilla length average 22.1 (18-27). One-half scales at mid- 
body average 27.2 (25-30), loreals 5.6 (4-8), temporals 13.1 (11-15), 
and fourth toe subdigital tricarinate scales 24.1 (18-28). The pari- 
etals are in contact slightly more often (58 per cent) than not, and 
the semicircles are more often complete (58 per cent) than not. 
The median head scales vary from 4 to 6 (mode 4), the prefrontal 
row is usually composed of three scales and is complete. The 
number of frontoparietals varies from 5 to 7 (mode 7), and only 
one lizard has the frontoparietal series interrupted. 

The paratypic males are dorsally like the type; one has Zone 6 
white and very bold in contrast to the adjacent ones. All had orange 
in the groin, in Zone 7, and on the hindlimbs. The venter is usu- 
ally immaculate cream, although a small subadult male (snout-vent 



Schwartz: New Lizards from Cuba 219 

57) shows some indication of ventral spotting like the females. All 
paratypes have the throat pattern more distinct and complete than 
does the type. The two V's and the two pairs of longitudinal 
lines are well expressed on a clear and unclouded cream ground 
color, and in only two specimens is the paramedian pair of lines 
joined to the posterior V. Three males show some dark orange 
necking on the underside of the limbs and basal portion of the 
tail. The dark throat pattern merges with orange spots and dots 
on the chest. These spots continue laterally and posteriorly onto 
the sides of the abdomen and merge with the orange pigment of 
Zone 7. 

The females have contrasting dorsal longitudinal lines as do 
the males, although Zones 4 and 6 are duller than in the males. 
The dorsolateral and lateral fields (Zones 3 and 5) have a series 
of transverse or vertical black bars which obscure to a large extent 
the dark brown ground color of these fields. Zone 7 likewise has 
some black vertical barring. The dark throat pattern is just as 
clear-cut in females as males; the bellies are spotted with brown 
(laterally) to orange (centrally), and these ventral spots may be 
elongate to form rather ill-defined longitudinal dashes. The hind 
limbs are rather dull orange; there are no dark dots or dashes on 
the underside of the hind limbs or tail. 

Comparisons. L. s. ophiplacodes is by far the most brilliantly 
colored of the three Camagiieyan subspecies of stictigaster. In 
coloration it is readily distinguished from both lucianus and para- 
sphex by the orange groin and orange hind limbs. The throat 
pattern differs from that of parasphex in that it is bold and black 
on a clear cream ground rather than diffuse and gray on a gray- 
clouded yellow ground. From lucianus, the throat pattern of 
ophiplacodes differs in lacking the pre-V transverse dark dash. 
Ventrally, lucianus and ophiplacodes males are readily distinguished 
by the presence of dark dotting in the former. Females of lucianus 
and ophiplacodes are similar, although female ophiplacodes are 
more brilliantly colored than those of lucianus and have transverse 
or vertical black bars in the fields, a feature which is absent in 
female lucianus. 

Although male ophiplacodes from the type locality do not reach 
so large a size as do males of the other two eastern races, females 
exceed female lucianus and parasphex in snout-vent length. The 
discrepancy in male size is possibly due the small sample. Although 



220 Quarterly Journal of the Florida Academy of Sciences 

ophiplacodes averages higher in both dorsal crest counts, loreals, 
and temporals, the amount of overlap is very great. Only in mid- 
body scales is there a higher average (27.2) in ophiplacodes than in 
lucianus (23.8) and paraphex (22.9), with less overlap of the counts 
(21 to 27 in both lucianus and paraphex, 25 to 30 in ophiplacodes). 

Although comparisons between the Camagiiey races are more 
pertinent, these races may be compared with the western Cuba 
and Isla de Pinos races as well. Of the eastern races, only astictus 
has the parietals more often not in contact as does parasphex. 
These two races can be differentiated by color and pattern; astictus 
has the sides red with green dots, a feature not found in para- 
phex (in fact, all Camagiiey races lack green in the pattern com- 
pletely). Both races have immaculate bellies, but the orange to 
red lateral dots of parasphex are not found in astictus. The dorsal 
crest scales in occiput to vent in parasphex average higher than 
those of all western forms except s. stictigaster, and the dorsal crest 
scales in occiput-axilla average lower than all races except sierrae. 

L. s. ophiplacodes reaches a larger size than all the eastern 
races, although it is approached by L. s. stictigaster. Dorsal crest 
scales in occiput-vent average greater in ophiplacodes than in any 
other subspecies. Only nominate stictigaster has the parietal con- 
tact relationship (slightly more often in contact) the same as in 
ophiplacodes. Detailed comparison of coloration and pattern is 
unnecessary; the eastern race is so much more brightly colored 
with its orange hind legs and bold throat pattern than any of the 
western subspecies that it is at once distinguishable on these fea- 
tures alone, without resort to scale counts and measurements. 

Etymology. From Greek ophis (serpent) and plakodes (plain- 
dweller), in reference to the serpentine savannas of central Cam- 
agiiey. 

Discussion 

With the occurrence of L. stictigaster now well established in 
Camagiiey Province, there remain three anomalous specimens from 
Oriente. Two of these in the Museo Poey, Universidad de la Ha- 
bana, are no longer available to me. I commented (1959, p. 110) 
that these "two Bayamo specimens . . . are even more peculiar; 
they are distinctly lined dorsally, have well defined zones 4 and 6, 
and lack a postorbital spot. The male is dotted ventrally and has 
dots in the lateral fields; both individuals have heavily marked 



Schwartz: New Lizards from Cuba 221 

throats, . . The dorsal crest scales are 51 and 56". Since the above 
was written I have examined another large adult male (MBZH 
130) also from Bayamo, Oriente Province, and collected by Charles 
T. Ramsden. The specimen is presently before me, and it agrees 
in detail with the two Bayamo specimens mentioned above. The 
dorsal crest scales are 54 between occiput and vent, giving the 
range of 51 to 56 for the three known extant Oriente specimens. 
The throat pattern is bold on an unclouded ground, the brown lat- 
eral fields have prominent dark brown dashes, and the belly is 
marked with dark brown longitudinal dashes. The snout-vent 
length is 87; this lizard is thus the largest specimen of L. stictigaster 
I have examined. Of the two Museo Poey specimens, the male has 
a snout-vent length of 86, the female a snout-vent length of 78; 
all three Bayamo specimens are thus very large. 

I have no doubt that these three lizards represent yet another 
subspecies of L. stictigaster from the Bayamo region, character- 
ized by large size, high number of dorsal crest scales, and distinc- 
tive throat and body pattern. However, since there are only three 
specimens available, and since two of these now are not before me, 
there seems little justification in describing this population. It is 
sufficient for the moment to acknowledge the occurrence of L. 
stictigaster in western Oriente and hope that at some future date 
more Oriente material will become available. 

The apparently split distribution of L. stictigaster, with one 
group of subspecies on the Isla de Pinos and in Pinar del Rio, and 
another group in Camaguey and Oriente, is puzzling. Although 
L. stictigaster does not occur with L. cubensis anywhere in the 
ranges of the two species, these two forms have peculiarly comple- 
mentary distributions, for example, on the Isla de Pinos (two sub- 
species of stictigaster separated by a race of cubensis), and in north- 
ern Camaguey (cubensis in the Sierra de Cubitas and associated 
mesic foothill areas, stictigaster both north and south of this moun- 
tain range on the xeric coast and the xeric savannas). In Pinar del 
Rio stictigaster is sympatric with L. carinatus, as it is widely and 
in detail elsewhere. In some areas (i.e., Playa Santa Lucia, Cama- 
guey) stictigaster also occurs with L. macropus. The vast interior 
of Cuba, however, is unoccupied by Leiocephalus except for L. 
cubensis. It seems hardly likely, despite the fact that cubensis 
and stictigaster do not occupy the same areas, that the latter species 
is unable to survive somewhere between central Pinar del Rio 



222 Quarterly Journal of the Florida Academy of Sciences 

and east-central Camaguey. Such a hiatus seems more probably 
to be a result of inadequate collecting in the intervening area rather 
than a reflection of a really disjunct distribution. 

Literature Cited 

Schwartz, Albert. 1959. Variation in lizards of the Leiocephalus cuben- 
sis complex in Cuba and the Isla de Pinos. Bull. Florida State Mus., 
vol. 4, no. 4, pp. 97-143, figs. 1-10. 

. 1960. A new subspecies of Leiocephalus stictigaster Schwartz from 

central Cuba. Proc. Biol. Soc. Washington, vol. 73, pp. 103-106. 

10000 SW 84 Street, Miami, Florida. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



NOTES ON FOSSIL TURKEYS 

Pierce Brodkorb 

These notes document certain changes in the classification of 
the turkeys adopted in the second part of the Catalogue of Fossil 
Birds. 

1. Paracrax Brodkorb 

Poracrax Brodkorb, 1964, p. 303 (type by original designation Meleagris antiqua 
Marsh). 

Both Shufeldt (1913) and Howard (1963) suggested that Mele- 
agris antiqua Marsh (1871) is not a turkey. Study of the holotype, 
the distal end of a right humerus, Yale Peabody Museum no. 537, 
confirms this view and indicates that it is referable to the family 
Cracidae. The ectepicondyle is low, lying below the level of the 
proximal end of the external condyle, with the tubercles protrud- 
ing laterally. The pit for the palmar branch of flexor carpi ulnaris 
is very deep. The external condyle has its distal end elevated above 
the internal condyle, as in Crax. The entepicondyle is broad in 
medial view, with the anconal margin forming an angle of about 
45 degrees, not compressed into a point as in the turkeys. 

2. Agriocharis progenes, new species 
Agriocharis progenes Brodkorb, 1964, p. 324 (nomen nudum). 

Holotype. Distal part of right tarsometatarsus with spur core 
(pi. 1, upper figs.), University of Michigan Museum of Paleontology 
no. 31034. From Rexroad Formation, uppermost Pliocene, at lo- 
cality 3, Rexroad ranch, Meade County, Kansas. Collected by 
Claude W. Hibbard and party, summer 1953. 

Referred material. All specimens are from Rexroad locality 3 
and are preserved in the University of Michigan. 

Premaxilla, no. 31052. Dentary, no. 47783. Portion of sternal 
carina, no. 31029. Left coracoid, lacking distal end (male), no. 
29040; fragmentary upper part of right coracoid (male), no. 48193. 
Right ulnare (female), no. 48190. Complete left carpometacarpus 
(male), no. 29041; distal end of two right carpometacarpi (females), 
nos. 48188, 48194. Right femur, lacking distal end (male), no. 
45912. Distal end of two left tibiotarsi (females), nos. 45970, 48191. 









mm 



Brodkorb: Fossil Turkeys 225 

Distal end of right tarsometatarsals (female), no. 48189. At least 
one male and two females are included. 

Tentatively referred material. Proximal part of right scapula, 
no. 45930; proximal part of juvenile left scapula, no. 45965. Both 
are small and represent females if correctly referred. 

Diagnosis. Tarsometatarsus with spur core situated low and 
medially directed at angle of about 50 degrees to acrotarsial face 
of bone (core low and at about 45 degrees in living A. ocellata of 
Yucatan; in A. crassipes, from the Upper Pleistocene of Nuevo Leon, 
core slightly higher but angle 39 degrees; in A. leopoldi, from the 
Lower Pleistocene of Texas, core slightly higher and angle greater, 
53-58.5 degrees; in Meleagris and Parapavo core elevated and di- 
rected more to rear, at angle of about 60-80 degrees). Facet for 
hind toe low (rather low in A. ocellata; elevated in Meleagris). Lat- 
eral distal foramen low (as in A. ocellata; apparently slightly higher 
in A. leopoldi; high in Meleagris and Parapavo). Inner distal fora- 
men small but well developed in both specimens. Trochleae more 
divergent and intertrochlear spaces wider than in other turkeys. 
Inner trochlea relatively narrow (more as in Meleagris; wider in 
A. ocellata and A. leopoldi). Inner trochlea less deflected to rear 
than in other turkeys. 

Height of middle of spur core above tip of middle trochlea, 
48.8; least width of shaft, 8.1, 2 6.8; width through trochleae, 7.5, 
2 6.3; width of middle trochlea, 7.5, 2 6.3 mm. 

A male tarsometatarsus illustrated by Wetmore (1924), from 
the Upper Pliocene at the Gum ranch, near Benson, Arizona, is 
similar in size and position of the spur core and is apparently refer- 
able to A. progenes. 

Both A. leopoldi (A. H. Miller and Bowman, 1956) and A. cras- 
sipes (L. Miller, 1940) have the spur core low on the shaft and at 
an angle of less than 60 degrees, characters that require their re- 
moval from the genus Meleagris, in which they were described. 

Premaxilla and dentary relatively short, wide, and only slightly 
vaulted (as in A. ocellata; longer, narrower, and more vaulted in 
Meleagris). The lower surface of the premaxilla has a wide trans- 



Plate 1. Fossil turkeys. Upper figures: Agriocharis progenes n. sp., 
hole-type tarsometatarsus, Rexroad, Kansas. Lower left: A. progenes, referred 
femur, Rexroad, Kansas. Lower right: Agriocharis anza Howard, referred 
femur, Rattlesnake Point, Texas. 



226 Quarterly Journal of the Florida Academy of Sciences 

verse bridge anterior to the palatine vacuity, leaving a large fora- 
men anterior to each choana. One specimen of A. ocellata ap- 
proaches this condition by having a small prong extending laterally 
from the median area of the elongate choanae. Premaxillary 
length, from tip to nostril, 13.8; width at nostrils (restored), 13.4 
mm. Ventral length of gonys, without shelf, 8.5; width of dentary 
at posterior end of gonys, 9.0; depth at the same point, 3.0 mm. 

Sternum too fragmentary for description. 

Coracoid with upper end much more strongly inflected than 
in other turkeys, at 75 degrees to axis of shaft (not over 60 degrees 
in A. ocellata and Meleagris). Head raised well above inner sur- 
face of neck, with bounding groove extending across inner side of 
bone (as in A. ocellata; in Meleagris neck merging gently with head, 
with groove incipient and present only near lip). Scapular facet 
obliquely elliptical (as in A. ocellata and Meleagris; rounded in 
Parapavo); its lower margin obsolete near procoracoid process (mar- 
gin raised throughout in the others). Inner posterior intermuscular 
line slightly curved away from inner edge of shaft (more than in 
A. ocellata, less than in Meleagris and Parapavo). Outer posterior 
intermuscular line more curved away from outer edge of shaft 
than in the others. Size near that of male A. ocellata, but shaft 
relatively wider than in that or other species. Length to pneumatic 
foramen, 65.0; head through scapular facet, 31.0; width of head, 
10.8; least width of shaft, 10.1 mm. 

The two tentatively referred scapulae differ from all known 
turkeys in lacking the pneumatic foramen. Wetmore (1944) like- 
wise recorded (as Meleagrididae, sp. ?) a non-foraminate scapula 
from the same locality. 

Ulnare agrees with A. ocellata in having ulnar base short, 
high, and wide (in Meleagris ulnar base long, low, and narrow). 
Size smaller than in either living turkey. Length of ulnar base, 
8.5; height of base, 4.9; width of ulnar facet, 4.4; height through 
uncinate process, 10.4 mm. 

Carpometacarpus with edge of inner trochlea deeply notched 
proximally by Ligamentum internum ossi carpi ulnaris et meta- 
carpi (as in Meleagris; notch shallower in A. ocellata). Carpal fos- 
sae relatively shallow (as in A. ocellata; somewhat deeper in Mele- 
agris). Intermetacarpal tubercle well proximal (located more dis- 
tally in Meleagris). Shaft of metacarpal II wide (as in A. ocellata; 
narrower in Meleagris). Facet for digit III longer than in living 



Brodkorb: Fossil Turkeys 227 

turkeys. Length, 66.6; height through metacarpal I, 20.0; width 
through trochleae, 9.5; least width of shaft, 8.0, 6.3, 5.8; width of 
facet for digit II, 6.1, 4.9; protrusion of metacarpal III beyond 
knob of metacarpal II, 3.8 mm. The large complete specimen re- 
sembles male A. ocellata in size; the two fragmentary ones are 
somewhat smaller than the female of that species. 

Femur (pi. 1, lower left) agrees with that of A. ocellata in hav- 
ing groove for Ligamentum capsulare femoris shallow and only 
slightly notching lesser trochanter (groove deep and lesser tro- 
chanter strongly notched in Meleagris and Parapavo). Posterior 
intermuscular lines fused along middle third of their length (as 
usual in Agriocharis and Parapavo; in Meleagris the lines are usu- 
ally unfused, although the character is variable). Size near that 
of male A. ocellata, but shaft narrower distally. Proximal width, 
25.8+; width below head, 20.0+ ; least width of shaft, 9.2; least 
depth of shaft, 8.4 mm. 

Tibiotarsi of the three genera of turkeys seem indistinguish- 
able except on size. The two fossils are small, resembling female 
A. ocellata, and thus smaller than Meleagris and Parapavo. Distal 
width (48191), 14.5; depth of internal condyle (48191), 15.2; depth 
of external condyle (45970), 13.0 mm. 

Wetmore (1944) reported a tibiotarsus from Rexroad locality 
3 as Meleagris gallopavo, noting that it was small. In the light of 
our present knowledge of the Rexroad turkey, this specimen may 
be referred to A. progenes, and the record of the living wild tur- 
key should be deleted from the Pliocene. 

Etymology. Greek progenes (born in olden times), referring 
to the fact that this is the oldest turkey known. 

3. Agriocharis anza Howard 

Agriocharis sp., Hibbard, 1960, p. 20 (Knox County, Texas). 

Agriocharis anza Howard, 1963, p. 19, pi. 3 (Middle Pleistocene, Palm Springs 
Formation, Arroyo Tapiado, California; type right humerus and associ- 
ated fragments of left humerus, sternum, sacrum, and ulna, Los Angeles 
County, Mus.). 

Referred specimen. Right femur, lacking distal end, Univ. 
Mich. Mus. Paleo. no. 39387, from Middle Pleistocene, Seymour 
Formation, at Rattlesnake Point, north side of South Fork of 
Wichita River, 4 miles south and x k mile east of Gilliland, Knox 



228 Quarterly Journal of the Florida Academy of Sciences 

County, Texas; collected by Walter W. Dalquest, March 6, 1956 
(pi. 1, lower right). 

Description. Lesser trochanter unnotched and groove for Lig- 
amentum capsulare femoris obsolete; posterior intermuscular lines 
fused along middle third of their length; medial border of shaft 
flaring broadly before meeting head of femur. 

Size near that of A. progrenes and male A. ocellata, but shaft 
wide proximally. Proximal width (head eroded), 25.5+; width 
below head, 21.4; least width of shaft, 10.4; least depth of shaft, 
9.1; width of head (eroded), 9.2+ mm. 

Since absence of comparable elements prevents direct compari- 
son, the Knox County specimen is referred to A. anza because of 
agreement in geologic horizon, general size, and marked expansion 
of the shaft of the femur (shaft of humerus expanded in type of 
A. anza). 

Acknowledgments. I am indebted to Elwyn L. Simons and 
James A. Hopson for permission to examine Marsh's types and to 
Claude W. Hibbard for the opportunity to study the fossils from 
Kansas and Texas. This study was aided by the National Science 
Foundation through grant number GB-1686. 

Literature Cited 

Brodkorb, Pierce. 1964. Catalogue of fossil birds: part 2 (Anseriformes 
through Galliformes. Bull. Florida State Mus., vol. 8, no. 3, pp. 195- 
335. 

Hibbard, Claude W. 1960. An interpretation of Pliocene and Pleistocene 
climates of North America. Rept. Michigan Acad. Arts, Sci., Letters 
for 1959-60, pp. 5-30, text-figs. 1-2, pi. 1. 

Howard, Hildegarde. 1963. Fossil birds from the Anza-Borrego desert. 
Los Angeles County Mus., Contr. in Sci., no. 73, pp. 1-33, pis. 1-3. 

Marsh, O. C. 1871. Notice of some new fossil mammals and birds, from 
the Tertiary formation of the west. Amer. Jour. Sci., ser. 3, vol. 2, pp. 
120-127. 

Miller, Alden H., and Robert I. Bowman. 1956. Fossil birds of the late 
Pliocene of Cita Canyon, Texas. Wilson Bull., vol. 68, no. 1, pp. 
38-46, fig. 1. 

Miller, Loye. 1940. A new Pleistocene turkey from Mexico. Condor, vol. 
42, no. 3, pp. 154-156, figs. 44-45. 



Brodkorb: Fossil Turkeys 229 

Shufeldt, R. W. 1913. Contributions to avian paleontology. Auk, vol. 30, 
no. 1, pp. 29-39, pi. 3. 

Wetmore, Alexander. 1924. Fossil birds from southeastern Arizona. Proc. 
U. S. Nat. Mus., vol. 64, art. 5, pp. 1-18, figs. 1-9. 

. 1944. Remains of birds from the Rexroad fauna of the Upper Plio- 
cene of Kansas. Univ. Kansas Sci. Bull., vol. 30, pt. 1, no. 9, pp. 
89-105, figs. 1-19. 

Department of Biology, University of Florida, Gainesville, 
Florida. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



OSTEOLOGY OF GALLINACEOUS BIRDS 
J. Alan Holm an 

The order Galliformes comprises a large group of almost world- 
wide distribution, yet its families are quite similar in basic morph- 
ology, and interfamilial hybrids occur. These factors have made 
a natural classification difficult, and at the present time several 
systems are in use. 

The classification generally followed in the United States is 
that of Wetmore (1960). He divides the order Galliformes into 
two suborders, the Opisthocomi which includes the hoatzins of 
the family Opisthocomidae, and the suborder Galli for the remain- 
ing groups. The suborder Galli is divided into two superfamilies, 
the Cracoidea, and the Phasianoidea. The superfamily Cracoidea 
includes the families Megapodiidae (mound-builders), and Cracidae 
(curassows). The superfamily Phasianoidea includes the families 
Tetraonidae (grouse), Phasianidae (pheasants, quails, peacocks), 
Numididae (guineafowl), and Meleagrididae (turkeys). 

Ridgway and Friedmann (1946) follow the classification of Wet- 
more, but further divide the Phasianidae into three subfamilies, 
the Odontophorinae (New World quails), Perdicinae (Old World 
quails), and Phasianinae (pheasants, junglefowls, peacocks). 

Other interpretations are given by Stresemann (1959), who treats 
the Galli and Opisthocomi of Wetmore as separate orders; and by 
Sibley (1960), who considers the order Galliformes to be composed 
of three families, the Megapodiidae, the Phasianidae (with sub- 
families Phasianinae, Meleagridinae, Numidinae, Tetraoninae, and 
Cracinae), and the Opisthocomidae. 

Mayr and Amadon (1951) consider the order Galli (Galliformes 
of Wetmore) to be composed of five families, Megapodiidae, Craci- 
dae, Phasianidae (with subfamilies Phasianinae, Numidinae, and 
Tetraoninae), Meleagrididae, and Opisthocomidae. 

Brodkorb (1964) recognizes no suborders or superfamilies in the 
Galliformes and recognizes five families, Cracidae (with living 
subfamilies Cracinae and Penelopinae), Opisthocomidae, Megapo- 
diidae, Numididae, and Phasianidae (with subfamilies Odonto- 
phorinae, Phasianinae, Tetraoninae, and Meleagrinae). 

Several recent studies utilizing new fossil material or employ- 



Holman: Osteology of Galliformes 231 

ing new approaches and techniques have attempted to evaluate 
the relationships among the Galliformes. These include papers by 
Tordoff and Macdonald (1957), early Oligocene fossils; Taibel 
(1961), physio-ethological characters; Hudson, Lanzollotti, and Ed- 
wards (1959), pelvic limb musculature; Sibley (1960), egg-white 
protein electrophoresis; and Mainardi (1960, 1963), Mainardi and 
Guerra (1959), and Mainardi and Taibel (1962 a and b), immuno- 
logical studies. 

During a recently published study of postcranial osteology of 
fossil and living New World quails (Holman, 1961), skeletons of 
representatives of the families of galliform birds were examined in 
order to ascertain the status of the New World quails. It then 
became evident that in many cases the relationships between 
gallinaceous groups were reflected by the postcranial skeleton. 
With the study of additional material the present paper has grown 
out of the earlier work. 

The classification of Wetmore as modified by Ridgway and 
Friedmann will be followed in the present paper through the sec- 
tions on comparative osteology and evolutionary trends. 

Acknowledgments 

I wish to most gratefully acknowledge the help of Pierce Brod- 
korb who supervised the problem of which this paper is a part. 
I also wish to thank others who have critically read the manu- 
script: Oliver L. Austin, Jr., Eugene Bovee, Harold K. Brooks, 
James N. Layne, E. Lowe Pierce, and William J. Riemer. 

William Quinn of Illinois State University made the drawings. 

Materials and Methods 

The skeletal collection of Pierce Brodkorb at the University of 
Florida and that of Robert D. Weigel at Illinois State University 
were supplemented by material borrowed from the United States 
National Museum (through Herbert Friedmann), University of Cali- 
fornia Museum of Vertebrate Zoology (through Frank A. Pitelka), 
University of Michigan Museum of Zoology (through H. B. Tor- 
doff), University of Kansas Museum of Natural History (through 
Richard F. Johnston), and Southern Illinois University Wildlife 
Research Collection (through W. D. Klimstra). 



232 Quarterly Journal of the Florida Academy of Sciences 

The number of specimens studied is listed below, with incom- 
plete skeletons given in parentheses. 

Megapodiidae: Alectura lathami 1, Leipoa ocellata 1, Macro- 
cephalon maleo 1, Megapodius freycinet 1. 

Cracidae: Crax globulosa 1, C. rubra 1, Mitu mitu 1, Ortalis 
vetula (4), Penelopina nigra (1). 

Tetraonidae: Bonasa umbeUus 6 (1), Canachites canadensis 
1 (2), Centrocercus urophasianus 4 (1), Dendragapus fuliginosus 
(1), D. obscurus 3 (1), Lagopus lagopus 4, L. leucurus 1, L. mutus 
2, Lyrurus tetrix 1 (3), Tetrao parvirostris 1, Tympanuchus cupido 
3 (1), T. pallidicinctus 2 (2), Pedioecetes phasianellus 2. 

Numididae: Acryllhim vulturinum 7, Nurnida meleagris 2. 

Meleagrididae: Meleagris gallopavo 8. 

Phasianidae, Phasianinae: Argusianus argus 1, Catreus wallichii 
1, Chrysolophus pictus 1, Galhts gallus 3, Gennaeus nycthemerus 1, 
Lophophorus impejanus 1, Prtuo cristatus 1, P. muticus 2, Phasianus 
colchicus 3. 

Phasianidae, Perdicinae: Alectoris graeca 2, A. ra/a, Coturnix 
coturnix 3 (1), Francolinns bicalcaratns (1), Perdix perdix 1 (1). 

Phasianidae, Odontophorinae: Callipepla squamata 4, Colinus 
leucopogon 4, C. nigrogularis (2), C. virginianus 99 (5), Cyrtonyx 
montezumae 3, Dactylortyx thoracicus 3, Dendrortyx leucophrys 1, 
Lophortyx californica 2 (1), L. douglasii 2, L. gambelii 2, Odonto- 
phorus gnjanensis 1 (1), O. guttatus 1, O. stellatus 1, Oreortyx picta 
4, Philortyx fasciatus 1, Rhynchoriyx cinctus 1. 

Opisthocomidae: Opisthocomus hoazin 2. 

Anatomical nomenclature in this paper follows that of Howard 
(1929). 

Comparative Osteology 

Characters found useful in the definition of galliform groups 
and those showing interrelationships are presented in this section. 
With one exception these are qualitative characters. A large num- 
ber of intermembral proportions were taken (many of these are in- 
cluded in the doctoral dissertation of J. Alan Holman at the Uni- 
versity of Florida Library, 1961), but most of these were of no 
value in the definition of group relationships. The skeletal ele- 
ments discussed are those most frequently found as fossils and 
with one exception are postcranial. Statements made in this sec- 
tion are based only on the specimens listed above. 



Holm an: Osteology of Galliformes 233 

Rostrum (Plate 3) 

Long, deep, strongly decurved (Cracidae); short, deep, strongly 
clecurved (Odontophorinae); long, shallow, slightly to moderately 
decurved (other Galliformes). 

Dorsal bony knob present (Mitu, Crax of Cracidae); bony knob 
absent (other Galliformes). 

Nasal fossae reduced (Opisthocomidae); fossae large (other Gal- 
liformes). 

Process of nasal bone projecting anteriorly from middle of pos- 
terior margin of nasal fossae (Macrocephalon of Megapodiidae; 
Opisthocomidae); without this condition (other Galliformes). 

Sternum (Plate 1) 

Manubrial spine ankylosed to furculum (Opisthocomidae); man- 
ubrial spine free anteriorly (other Galliformes). Manubrial spine 
with very large dorsal foramen (Cracidae); spine with moderately 
large dorsal foramen (Catreas, Phasianus, Argusianus of Phasiani- 
nae; Numididae); spine with slight pneumatic perforations dorsally 
(some Tympanuchus cupido, Lagopus mutus, Lyrurus tetrix, Bonasa 
umbellus of Tetraonidae); spine with obsolete dorsal foramen (some 
Pedioecetes of Tetraonidae); spine without dorsal foramina (other 
Galliformes). 

Anterior lateral processes absent (Opisthocomidae); anterior 
lateral processes present, short, broad, at right angle to long axis 
of sternum (Megapodiidae; Cracidae); anterior lateral processes 
present, short, broad, at about 45 degree angle to long axis of ster- 
num (Numididae); anterior lateral processes present, moderately 
short and wide, nearly parallel to long axis of sternum (Tetrao 
parvirostris of Tetraonidae; Pavo of Phasianinae); anterior lateral 
processes present, elongate and slender, nearly parallel to long 
axis of sternum (other Galliformes). 

Anterior sternal plate highly pneumatic (Megapodiidae; Craci- 
dae; Opisthocomidae); plate moderately pneumatic (Numididae; 
Meleagrididae); plate moderately or slightly pneumatic (Tetraoni- 
dae; Phasianinae); plate moderately, slightly or non-pneumatic 
(Odontophorinae); plate slightly or non-pneumatic (Perdicinae). 

Sternum with only one pair of short notches (Opisthocomidae); 
sternum with two pairs of notches (other Galliformes). Inner ster- 
nal notches short, thus posterior lateral and posterior medial proc- 



234 Quarterly Journal of the Florida Academy of Sciences 




Plate 1. Sternum of Galliformes. A, Crax globulosa; B, Acryllium vul- 
turinum; C, Phasianus colchicus; D, Opisthocomus hoazin. 



Holm an: Osteology of Galliformes 235 

esses arising independently from sternum (Megapodiidae; Craci- 
dae); inner sternal notches longer, thus posterior lateral and pos- 
terior medial processes arising from common base (other Galli- 
formes). 

Carina reduced in extent, but swollen, apex directed ventrally, 
posterior face excavated as elliptical concavity, without antero- 
lateral extending ridges (Opisthocomidae); carina well developed, 
narrow throughout, apex rotated anteriorly, without elliptical con- 
cavity, with anterolateral extending ridges (other Galliformes). 

Coracoid (Plate 3) 

Medial surface of head fused to furculum (Opisthocomidae); 
medial surface of head not fused to furculum, flattened (Meleagri- 
didae) or rounded (other Galliformes). 

Brachial tuberosity fused to furculum (Opisthocomidae); tuber- 
osity not fused to furculum, without overhanging ventral portion 
(Megapodiidae; Cracidae; Meleagrididae) or with overhanging 
ventral portion (other Galliformes). 

Dorsal intermuscular line not raised distally (Megapodiidae; 
Cracidae; most Phasianinae; Meleagrididae); line slightly raised 
distally (Gallus, Chrysolophus of Phasianinae; Numididae); line 
slightly or sharply raised distally (Tetraonidae); line sharply raised 
distally (Odontophorinae; Perdicinae; Opisthocomidae). 

Distal dorsal face without pneumatic fossa (most Megapodii- 
dae; Odontophorinae; Perdicinae; Numididae); face with large 
pneumatic fossa (Leipoa of Megapodiidae; other Galliformes). 

Sterno-coracoidal process without terminal knob (Leipoa of 
Megapodiidae; Cracidae; Dendrortyx of Odontophorinae; Opistho- 
comidae); process ending in obsolete terminal knob (Philortyx of 
Odontophorinae; Phasianinae; Numididae; Meleagrididae); process 
ending in well developed terminal knob (other Galliformes). 

Scapula (Plate 2) 

Ventral base of glenoid facet with large pneumatic fossa (most 
Megapodiidae; Cracidae); facet with small pneumatic fossa (Argusi- 
anus of Phasianinae; some Opisthocomidae); facet without pneu- 
matic fossa (Megapodius of Megapodiidae; other Galliformes). 

Area mediad to glenoid facet in dorsal aspect with depression 
(Odontophorinae; Perdicinae); area with slight depression (Mega- 



236 Quarterly Journal of the Florida Academy of Sciences 

podius of Megapodiidae; Crax of Cracidae; Lophophorus, Phasi- 
anus, Chrysolophus of Phasianinae; Acryllium of Numididae); area 
without depression (other Galliformes). 

Bridge between acromion process and glenoid facet with dis- 
tinct pneumatic fossa (Tetraonidae; Pavo of Phasianinae); bridge 
moderately pneumatic (Opisthocomidae); bridge slightly pneumatic 
(Argusianus of Phasianinae); bridge non-pneumatic (other Galli- 
formes). 

Acromion process straight (Leipoa of Cracidae; Tetraonidae; 
Meleagrididae; Opisthocomidae); process deflected (other Galli- 
formes). 

Dorsal base of shaft with pneumatic fossa (Meleagrididae); base 
non-pneumatic (other Galliformes). 

Blade very elongate (Odontophorinae; Coturnix of Perdicinae); 
blade moderately elongate (Alectoris, Perdix of Perdicinae); blade 
moderately short and wide (Megapodius, Leipoa of Megapodiidae; 
Ortalis of Cracidae; Tetraonidae; most Phasianinae); blade very 
short and wide (Alectura, Macrocephalon of Megapodiidae; Pavo 
of Phasianinae; other Galliformes). Apex of blade without ter- 
minal expansion (Alectura, Macrocephalon of Megapodiidae; Craci- 
dae; Pavo of Phasianinae; Acryllium of Numididae; Opisthocomi- 
dae); apex terminally expanded (other Galliformes). 

Humerus (Plate 2) 

Pneumatic fossa very small (Megapodiidae; Cracidae; Opistho- 
comidae); fossa small, but somewhat larger than in preceeding 
groups (Numididae; Meleagrididae); fossa moderately enlarged 
(Tetraonidae; Perdix of Perdicinae; Phasianinae); fossa much en- 
larged (Odontophorinae; most Perdicinae), 

Internal anconal border of bicipital crest narrow (Odontophori- 
nae; most Perdicinae); border very wide (Megapodiidae); border 
moderately wide (Perdix of Perdicinae; other Galliformes). 

With inner shelf extending from medial bar to internal bicip- 
ital surface absent (most Odontophorinae, Alectoris, Coturnix of 
Perdicinae); inner shelf present, but incomplete (Megapodius of 
Megapodiidae; Tetraonidae; Odontophorus gujanensis of Odonto- 
phorinae; Opisthocomidae); inner shelf present, complete (other 
Galliformes). 

Fossa II (Ashley, 1941) absent (Megapodiidae; Cracidae; Opis- 



Holm an: Osteology of Galliformes 



237 




Plate 2. Osteology of Galliformes. Upper row: Pelvis of A, Phasianus 
colchicus; B, Colinus virginianus. 

Middle row: Dorsal view of scapula of A, Mitu mitu; B, Numida mele- 
agris; C, Meleagris gallopavo; D, Phasianus colchicus; E, Colinus virginianus. 

Lower row: Anconal view of humerus of A, Crax globulosa; B, Numida 
meleagris; C, Meleagris gallopavo; D, Phasianus colchicus; E, Colinus virgini- 
anus; F, Opisthocomus hoazin. 



238 Quarterly Journal of the Florida Academy of Sciences 

thocomidae); fossa II well developed (most Odontophorinae; Co- 
turnix of Perdicinae); fossa II very weakly developed (Dendrortyx 
and Odontophorus of Odontophorinae; other Galliformes). 

Bicipital crest with lateral margin truncated (Alectura of Meg- 
apodiidae; Mitu of Cracidae; Numididae; Opisthocomidae); margin 
rounded (other Galliformes). 

Median crest very strongly developed, knoblike (Opisthocomi- 
dae); crest only moderately developed, flattened (other Galli- 
formes). 

Deltoid crest low on shaft, apex well below level of pneumatic 
fossa and bicipital crest, rotated anconally so that apex visible in 
anconal view (Opisthocomidae); crest high on shaft, apex at level 
of middle of pneumatic fossa and bicipital crest, rotated palmarly 
so that apex not visible in anconal view (other Galliformes). 

Ulna 

External cotyla weakly developed (Meleagrididae); cotyla mod- 
erately developed (Acryllium of Numididae); cotyla well developed 
(other Galliformes). Margin of external cotyla flattened (Alectura 
of Megapodiidae; Crax globidosa of Cracidae; Dendrortyx, Odon- 
tophorus guttatus of Odontophorinae; Argusianus of Phasianinae; 
Meleagrididae); margin rounded (other Galliformes). 

Carpometacarpus (Plate 3) 

Pisiform process at level of ligamental attachment (most Meg- 
apodiidae; Cracidae; Tetraonidae; most Perdicinae; Acryllium of 
Numididae; some Opisthocomidae); process produced beyond level 
of ligamental attachment (Alectura of Megapodiidae; Coturnix of 
Perdicinae; other Galliformes). 

Intermetacarpal process absent (most Megapodiidae; Numidi- 
dae, Opisthocomidae); process represented by a minute point 
(Alectura of Megapodiidae; Cracidae; Pavo of Phasianinae); proc- 
ess weakly developed, not extending to level of metacarpal III 
(some individuals of Crytonyx of Odontophorinae; Coturnix of Per- 
dicinae; Gennaeus, Catreus of Phasianinae); process well developed, 
extending to metacarpal III (other Galliformes). 

Carpal trochlea with external rim continuing distad beyond 
ligamental notch (Megapodiidae; Cracidae; Opisthocomidae); with 
external rim ending at ligamental notch (other Galliformes). 



Holman: Osteology of Galliformes 



239 




240 Quarterly Journal of the Florida Academy of Sciences 

Metacarpal III about as wide as metacarpal II (Opisthocomi- 
dae); III much narrower than II (other Galliformes). 

Pelvis (Plate 2) 

Pelvis very wide and shallow (Tetraonidae); relatively narrow 
and deep (other Galliformes). 

Pectineal process represented by only a minute point (Mega- 
podiindae; Opisthocomidae); process well developed, long and nar- 
row (Perdicinae; most Phasianinae); process small, short (Pavo 
cristatus of Phasianinae; other Galliformes). Pectineal process with 
pneumatic foramen on anterior border (Leipoa, Alectura of Mega- 
podiidae; Cracidae); process without pneumatic foramina (Meg- 
apodius, Macrocephalon of Megapodiidae; most Odontophorinae; 
Perdicinae; Numididae); process with foramen or foramina on 
medial face (Dendrortyx of Odontophorinae; other Galliformes). 

Entire renal depression highly pneumatic (most Megapodii- 
dae); depression with a moderate number of pneumatic foramina 
(Megapodius of Megapodiidae); anterior renal depression with 
large pneumatic fossa (Cracidae Acryllhim of Numididae); ante- 
rior depression with moderately large pneumatic fossa (Numida 
of Numididae); anterior renal depression with a few small pneu- 
matic foramina (Bonasa of Tetraonidae; Lophophorus, Gallm, Ca- 
treus, Argusianus, Pavo of Phasianinae); renal depression non-pneu- 
matic (other Galliformes). 

Renal bar slender (Ortalis of Cracidae; Dendrortyx, Philortyx, 
Oreortyx, Callipepla, Colinus, Lophortyx of Odontophorinae; Co- 
turnix of Perdicinae); bar broad (other Galliformes). 

Roof formed by dorsal extensions of preacetabular ilia over 
synsacral vertebrae present (Opisthocomidae); preacetabular ilial 
roof absent (other Galliformes). 



Plate 3. Osteology of Galliformes. Upper left: Rostrum of A, Coturnix 
coturnix; B, Colinus virginianus. 

Second and third rows: Dorsal and ventral views of coracoid of A and E, 
Crax globulosa; B and F, Meleagris gallopavo; C and G, Phasianus colchicus; 
D and H, Colinus virginianus. 

Bottom row: External view of carpometacarpus of A, Crax globulosa; 
B, Meleagris gallopavo. 

Upper right: Anterior view of femur of A, Crax globulosa; B, Phasianus 
colchicus. 

Lower right: Anterior view of tarsometatarsus of A, Crax globulosa; B, 
Phasianus colchicus. 



Holman: Osteology of Galliformes 241 

Femur (Plate 3) 

Head flattened, dorsal border at level of iliac facet (Meleagridi- 
dae); head swollen, dorsal border above level of iliac facet (other 
Galliformes). 

Dorsal crest of trochanter very high (Megapodiidae; some Mele- 
agrididae); crest low (Cracidae; Rhynchortyx, Cyrtonyx, Odonto- 
phorus, Colinus of Odontophorinae; Opisthocomidae); crest mod- 
erately high (other Galliformes). 

Area mediad to anterior border of trochanter with pneumatic 
fossa (Alectura, Leipoa of Megapodiidae; Tetraonidae; most Phasi- 
aninae); area laterad to posterior aspect of head with pneumatic 
fossa (Argusianus of Phasianinae); both areas non-pneumatic (Lo- 
phophorus, Gallus of Phasianinae; other Galliformes). 

Round ligament attachment deeply excavated (Opisthocomidae); 
attachment shallow (other Galliformes). 

Tibiotarsus 

Inner cnemial crest usually arising from shaft well above level 
of outer cnemial crest (Meleagrididae); inner cnemial crest arising 
at level of outer cnemial crest (most Perdicinae; Numida of Numidi- 
dae); inner cnemial crest arising from shaft well below level of 
outer cnemial crest (Coturnix of Perdicinae; other Galliformes). 

Outer cnemial crest weakly developed (Opisthocomidae); outer 
crest strongly developed (other Galliformes). 

Tarsometatarsus (Plate 3) 

Hypotarsus with two roofed calcaneal canals (Lagopus lagopus 
I of 4, Bonasa umbellus 2 of 7, Pedioecetes phasianellus 1 of 2, 
Tympanuchus cupido 2 of 4 of Tetraonidae; most Odontophorinae); 
hypotarsus with one roofed calcaneal canal (Dendrortyx, Rhyn- 
chortyx, most Oreortyx of Odontophorinae; other Galliformes). 

Inner calcaneal ridge with distal extension running about one- 
third to three-fourths the way down shaft (most Tetraonidae; Alec- 
toris of Perdicinae; most Phasianinae; Meleagrididae); ridge with- 
out distal extension (Lagopus lagopus 2 of 4, Bonasa umbellus 4 
of 7, Centrocercus urophasianus 2 of 5 of Tetraonidae; Gallus of 
Phasianinae; other Galliformes). 



242 Quarterly Journal of the Florida Academy of Sciences 

Posterior shaft with spur core in male (Phasianinae; Meleagridi- 
dae); posterior shaft with rudimentary spur core in male (Alectoris 
of Perdicinae); posterior shaft without spur core (other Galliformes). 

Trochleae short, metacarpal III only slightly extending below 
level of metacarpals II and IV (Opisthocomidae); trochleae mod- 
erately long, metacarpal III extending well below level of meta- 
carpals II and IV (other Galliformes). 

Trochlea for digit II at level of trochlea for digit IV (Mega- 
podius, Macrocephalon of Megapodiidae; Cracidae; Opisthocomi- 
dae); II slightly elevated above IV (Leipoa, Alectura of Megapodii- 
dae); II well above IV (other Galliformes). 

Intermembral Proportions 

Pelvis longer than sternum (Megapodiidae; Opisthocomidae); 
pelvis shorter than sternum (other Galliformes). 

EVOLUTIONARY TRENDS 

Most authors have considered the superfamily Cracoidea to 
represent the most primitive of the gallinaceous birds. The recent 
work of Hudson, Lanzillotti, and Edwards (1959) on the pelvic limb 
musculature in galliform birds tends to confirm this view. More- 
over, Mainardi and Taibel (1962b) go so far as to state there is evi- 
dence that all galliform birds have descended from an ancestral 
stock similar to the family Cracidae. These remarks are based on 
morphological and paleontological data. If indeed this family 
represents the most primitive living galliform group, then several 
trends in other galliform taxa toward modification of structures 
found in the Cracidae become evident. These trends are sum- 
marized below. 

Sternum 

(1) Increased excavation of sternum by notches. Cracidae, 
Megapodiidae, Opisthocomidae: sternum excavated by one or two 
pairs of short notches. Other Galliformes: sternum excavated by 
two pairs of long notches. 

(2) Reduction and loss of the dorsal foramen in the manubrial 
spine. Cracidae: foramen very large. Numididae: foramen mod- 
erately large. Phasianinae: foramen moderately large or absent. 
Tetraonidae: foramen obsolete or absent. Megapodiidae, Melea- 



Holman: Osteology of Galliformes 243 

grididae, Perdicinae, Odontophorinae: foramen absent. In the 
Opisthocomidae the fusion of the manubrial spine with the furcu- 
lum obscures this character. 

(3) Modification of the anterior lateral processes. Cracidae, 
Megapodiidae: processes short, broad, at right angle to long axis 
of sternum. Numididae: processes short, broad, at 45 degree angle 
to long axis of sternum. Meleagrididae, Phasianinae (except Pavo), 
Perdicinae, Tetraonidae (except Tetrao parvirostris), Odontophori- 
nae: processes long, narrow, parallel to long axis of sternum. In the 
Opisthocomidae the anterior lateral processes are missing. 

(4) Decreasing pneumaticity of the anterior sternal plate. Opis- 
thocomidae, Megapodiidae, Cracidae: plate highly pneumatic. 
Numididae, Meleagrididae: plate moderately pneumatic. Tetra- 
onidae, Phasianinae: plate moderately or slightly pneumatic. Odon- 
tophorinae: plate moderately, slightly, or non-pneumatic. Perdici- 
nae: plate slightly or non-pneumatic. 

Coracoid 

(5) Development of an overhanging ventral portion of the brach- 
ial tuberosity. Cracidae, Megapodiidae, Meleagrididae: overhang- 
ing ventral portion absent. Other families: overhanging ventral 
portion present. 

(6) Development of a sharply raised distal portion of the dorsal 
intermuscular line. Cracidae, Megapodiidae, Meleagrididae: line 
without sharply raised distal portion. Phasianinae: line usually 
without sharply raised distal portion, but may be slightly raised 
distally. Numididae, some Tetaronidae: line slightly raised dis- 
tally. Opisthocomidae, most Tetraonidae, Perdicinae, Odontophor- 
inae: line sharply raised distally. 

(7) Loss of the large fossa on the distal dorsal face. Opistho- 
comidae, Cracidae, some Megapodiidae, Meleagrididae, Phasiani- 
nae, Tetraonidae: large pneumatic fossa present. Numididae, 
Odontophorinae, Perdicinae, most Megapodiidae: pneumatic fossa 
absent. 

(8) Development of a terminal knob on the sterno-coracoidal 
process. Opisthocomidae, Cracidae, Leipoa of Megapodiidae, Den- 
drortyx of Odontophorinae: sterno-coracoidal process without ter- 
minal knob. Numididae, Meleagrididae, Phasianinae, Philortyx of 



244 Quarterly Journal of the Florida Academy of Sciences 

Odontophorinae: terminal knob quite small. Most Megapodiidae, 
Perdicinae, Tetraonidae, most Odontophorinae: terminal knob well 
developed. 

Scapula 

(9) Loss of pneumatic fossa in the ventral base of the glenoid 
facet. All Megapodiidae but Megapodius, Cracidae: large pneu- 
matic fossa present. Opisthocomidae, Argusianus of Phasianinae: 
small pneumatic fossa sometimes present. Other forms (pneumatic 
fossa absent. 

(10) General narrowing of the blade. Opisthocomidae, Craci- 
dae except Ortalis, some Megapodiidae, Numididae, Meleagrididae, 
Pavo of Phasianinae: blade very short and wide. Some Megapodii- 
dae, Ortalis of Cracidae, Tetraonidae, most Phasianinae: blade mod- 
erately short and wide. Most Perdicinae: blade moderately elon- 
gate. Perdix of Perdicinae, Odontophorinae: blade very elongate. 

(11) Development of terminal expansion on blade apex. Opis- 
thocomidae, Cracidae, some Megapodiidae, Acryllium of Numidi- 
dae, Pavo of Phasianinae: apex without terminal expansion. Other 
forms: apex terminally expanded. 

Humerus 

(12) Increasing pneumaticity of the proximal end, including en- 
largement of the pneumatic fossa, obliteration of its inner shelf, 
and development of a second fossa. Opisthocomidae, Cracidae, 
Megapodiidae: pneumatic fossa very small, with inner shelf extend- 
ing from medial bar to internal bicipital surface; the shelf is in- 
complete in the Opisthocomidae and Megapodius of the Megapodii- 
dae. Numididae, Meleagrididae: pneumatic fossa small, but some- 
what larger than in preceeding forms; complete inner shelf pres- 
ent. Phasianinae, Perdix of Perdicinae, Tetraonidae (pneumatic 
fossa moderately large; inner shelf present, but incomplete in 
Tetraonidae), most Perdicinae; Odontophorinae: pneumatic fossa 
much enlarged; inner shelf absent except in Odontophorus guttatus 
of Odontophorinae. 

Opisthocomidae, Cracidae, Megapodiidae: fossa II absent. 
Tetraonidae, Numididae, Meleagrididae, Phasianinae, Alectoris, 
Perdix of Perdicinae, Dendrortyx, Odontophorus of Odontophori- 
nae: fossa II very weakly developed. Coturnix of Perdicinae, most 
Odontophorinae: fossa II well developed. 



Holm an: Osteology of Galliformes 245 

Carpometacarpus 

(13) Development of a long intermetacarpal process. Opistho- 
comidae, Cracidae, Megapodiidae, Numididae, Pavo of Phasiani- 
nae: intermetacarpal process absent or represented by only a minute 
point. Gennaeus, Catreus of Phasianinae, Coturnix of Perdicinae, 
some individuals of Cyrtonyx of Odontophorinae: intermetacarpal 
process weakly developed. Remaining forms: intermetacarpal 
process well developed, extending to level of metacarpal III. 

(14) Modification of the external rim of the carpal trochlea. 
Opisthocomidae, Megapodiidae, Cracidae: external rim of carpal 
trochlea continues distad beyond ligamental notch. Other forms: 
external rim of carpal trochlea ends at ligamental notch. 

Pelvis 

(15) Decreasing pneumaticity of renal depression. Some Opis- 
thocomidae, Cracidae, Megapodiidae, Numididae: renal depression 
much or moderately perforated by pneumatic foramina, or with a 
pneumatic fossa. Bonasa of Tetraonidae, hophophorus, Gallus, 
Catreus Argusianus, Pavo of Phasianinae: anterior renal depression 
with a few small pneumatic foramina. Other forms: renal depres- 
sion non-pneumatic). 

Tarsometatarsus 

(16) Elevation of the trochlea for digit II. Opisthocomidae, 
Cracidae, Megapodius, Macrocephalon of Megapodiidae: trochlea 
for digit II at level of trochlea for digit IV. Leipoa, Alectura of 
Megapodiidae: trochlea for digit II slightly elevated above trochlea 
for digit IV. Other forms: trochlea for digit II elevated well above 
trochlea for digit IV. 

Among the above characters the cracid-like condition is shown 
(at least by some genera) 14 times in the Megapodiidae, 12 times 
in the Opisthocomidae, five times in the Numididae, five times in 
the Meleagrididae, twice in the Phasianinae, once in the Tetraoni- 
dae, and never in the Perdicinae and Odontophorinae. 

Discussion 

In many cases the families of gallinaceous birds as outlined by 
Wetmore (1960) and the phasianid subfamilies as outlined by Ridg- 
way and Friedmann (1946) are difficult to define on the basis of post- 



246 Quarterly Journal of the Florida Academy of Sciences 

cranial osteology. Many characters that will distinguish most 
genera of some groups from most genera of other groups break 
down in a few genera, in a single genus, or even in species and indi- 
viduals. Moreover, there are few characters that are unique at 
the familial and subfamilial level other than in the Opisthocomidae. 

Several osteological characters are shared by the families Craci- 
dae, Megapodiidae, and Opisthocomidae. Indeed, I believe there 
is not much doubt that these families are much more closely re- 
lated to each other than to the remaining gallinaceous birds. 

Among these remaining galliform groups there seems to be a 
tendency for some groups to depart more radically than others 
from the primitive cracid-like condition. 

Following is a tentative arrangement of gallinaceous families 
and subfamilies based on the rostrum and postcranial osteology. 

Order Galliformes 

Family Cracidae 
Family Megapodiidae 
Family Opisthocomidae 
Family Numididae 
Family Meleagrididae 
Family Phasianidae 

Subfamily Tetraoninae 

Subfamily Phasianinae 

Subfamily Odontophorinae 

Family Cracidae 

The closest osteological affinities of the Cracidae are with the 
Megapodiidae. The following characters will separate the two 
families: 

Sternum: less than twice as long as inner notch. Megapodii- 
dae, more than twice as long as inner notch. Manubrial spine with 
very large dorsal foramen. Megapodiidae, dorsal foramen absent. 

Ulna: external cotyla weakly developed. Megapodiidae, ex- 
ternal cotyla well developed. 

Femur: dorsal crest low. Megapodiidae, dorsal crest very high. 

The next closest affinities of the Cracidae are with the Opistho- 
comidae. 



Holman: Osteology of Galliformes 247 

Family Megapodiidae 

The closest osteological affinities are with the Cracidae, but 
many characters are shared with the Opisthocomidae. 

Family Opisthocomidae 

The hoatzin appears to be at once primitive and very highly 
specialized. Its closest osteological affinities are with the Cracidae 
and Megapodiidae, but it has many unique characters as follow: 

Rostrum: nasal fossae reduced. Other forms, nasal fossae large. 

Sternum: manubrial spine ankylosed to furculum, sternal plate 
lacking anterior lateral processes, only one pair of very short notch- 
es; carina reduced in extent, but swollen, apex directed ventrally, 
posterior face excavated as elliptical concavity, lacking anterolat- 
eral extending carinal ridges. Other forms, manubrial spine free 
anteriorly; sternal plate with anterior lateral processes and two 
pairs of notches; carina well developed and narrow throughout, 
its apex directed anteriorly, elliptical concavity lacking on its pos- 
terior face, carinal ridges extending anterolaterally. 

Coracoid: medial surface of head and brachial tuberosity fused 
to furculum. Other forms, medial surface of head and brachial 
tuberosity free. 

Humerus: medial crest strongly developed, knoblike; deltoid 
crest low on shaft, its apex well below level of pneumatic fossa and 
bicipital crest; rotaed anconally so apex visible in anconal view. 
Other forms, median crest moderately developed, flattened; del- 
toid crest high on shaft, its apex at level of middle of pneumatic 
fossa and bicipital crest; rotated palmarly so apex not visible in 
anconal view. 

Carpometacarpus: metacarpal III about as wide as metacarpal 
II. Other forms, metacarpal III much narrower than metacarpal II. 

Pelvis: roof formed by dorsal extensions of preacetabular ilium 
over synsacral vertebrae present. Other forms, roof absent. 

Femur: round ligament attachment much excavated. Other 
forms, round ligament attachment shallow. 

Tibiotarsus: outer cnemial crest weakly developed. Other 
forms, crest strongly developed. 

Tarsometatarsus: trochleae short, metacarpal III extends only 
slightly below metacarpals II and IV. Other forms, trochleae 



248 Quarterly Journal of the Florida Academy of Sciences 

longer, metacarpal III extends well below level of metacarpals 
II and IV. 

Family Numididae 

This family has some osteological characters shared by the Craci- 
dae but it shows more similarities to other more advanced galli- 
naceous groups. There is one unique character as follows: 

Sternum: anterior lateral processes short, broad, making angle 
of about 45 degrees to long axis of sternum. Other forms, anterior 
lateral processes when present either long and narrow and parallel 
to long axis of sternum, or short and broad and at right angle to 
long axis of sternum. 

Family Meleagrididae 

Although the turkeys have a few cracid-like characters, two 
are unique. 

Scapula: dorsal base of shaft with pneumatic fossa. Other 
forms, dorsal base of shaft without fossa. 

Femur: head flattened, its dorsal border at level of iliac facet. 
Other forms, head swollen, its dorsal border above level of facet. 

Family Phasianidae 

With the exception of the genus Pavo, this heterogeneous group 
shows more modifications from the cracid skeleton than any other 
gallinaceous family. There are no osteological characters unique 
to the Phasianidae as a whole, which thus must be defined by a 
combination of characters. 

Subfamily Tetraoninae 

The grouse appear to have closest skeletal affinities with the 
pheasants, junglefowls, peacocks, and Old and New World quails, 
and for this reason are placed as a subfamily of the Phasianidae. 
The very shallow and wide pelvis is a unique character of the 
Tetraoninae. 

Subfamily Phasianinae 

This subfamily includes both the Phasianinae and Perdicinae 
of Ridgway and Friedmann (1946). On one hand the Old World 
quails (Perdicinae of Ridgway and Friedmann) show some basic 
resemblances to the pheasants, but on the other hand some of the 



Holman: Osteology of Galliformes 249 

same trends in modification of skeletal structures as found in the 
New World quails, probably through parallel evolution in the Old 
and New World birds. 

The genus Pavo has certain skeletal similarities to both Numida 
and Acryllium of the Numididae, not shared by other birds of the 
subfamily Phasianinae. Some of these include the short, broad an- 
terior lateral processes of the sternum, the lack of a well defined 
intermetacarpal process of the carpometacarpus, and the lack of a 
well developed pectineal process of the pelvis in Pavo cristatus. 
Perhaps Pavo is a linking genus between the Phasianinae and the 
Numididae. Mainardi and Taibel (1962b) state "Pavo is immuno- 
logically more closely related to Numida than to any other phasi- 
anid". They also state "... they hybridize easily, have quite 
similar karotypes, and the same modality of moult of the rec- 
trices . . . ". 

Subfamily Odontophorinae 

In a previous paper (Holman, 1961) I considered the New World 
quails to represent a separate family, the Odontophoridae. At 
the present time I feel that it is best to retain the New World 
quails as a subfamily of the Phasianidae because of the characters 
that are shared with the grouse, pheasants, junglefowl, peacocks, 
and Old World quails. 

The Odontophorinae comprise the phasianid group that departs 
most radically from the cracid-like skeleton, although some parallel 
developments have taken place in the Old World quails of the 
subfamily Phasianinae. Characters that separate the Odontophori- 
nae from the Phasianinae are as follow: 

In the rostrum, (1) Odontophorinae, rostrum short, deep, strong- 
ly decurved; Phasianinae, rostrum long, shallow, slightly decurved. 

In the sternum, (1) Odontophorinae, manubrial spine without 
dorsal foramen; Phasianinae, manubrial spine often with dorsal 
foramen. 

In the coracoid, (1) Odontophorinae, dorsal intermuscular line 
sharply raised distally; Phasianinae, dorsal intermuscular line usu- 
ally not raised, or only slightly raised distally. (2) Odontophorinae, 
ventral intermuscular line terminating at tip of sterno-coracoidal 
process or occasionally in middle of distal border of sternal facet; 
Phasianinae, ventral intermuscular line often terminating near lat- 
eral end of sternal facet. (3) Odontophorinae, distal dorsal face 



250 Quarterly Journal of the Florida Academy of Sciences 

non-pneumatic; Phasianinae, distal dorsal face almost always with 
a large pneumatic fossa. (4) Odontophorinae, sterno-coracoidal 
process usually terminating in well developed terminal knob; 
Phasianinae, sterno-coracoidal process usually terminating in ob- 
solete terminal knob. 

In the scapula, (1) Odontophorinae, scapular blade very elon- 
gate, with its dorsal surface grooved throughout, and with its apex 
expanding terminally; Phasianinae, scapular blade with much vari- 
ation in shape, but never as in Odontophorinae. 

In the humerus, (1) Odontophorinae, pneumatic fossa much 
enlarged, with inner shelf in only one species of Odontophorus; 
Phasianinae, pneumatic fossa usually moderately large, usually 
with inner shelf extending from medial bar to internal bicipital 
surface. (2) Odontophorinae, fossa II usually well developed; Phas- 
ianinae, fossa II usually obsolete. 

In the pelvis, (1) Odontophorinae, pectineal process obsolete; 
Phasianinae, pectineal process well developed, long (Pavo cristatus 
is the single exception with the pectineal process obsolete, whereas 
it is rather well developed in Pavo muticus). (2) Odontophorinae, 
renal bar non-pneumatic; Phasianinae, renal bar often with pneu- 
matic foramen. 

In the tarsometatarsus, (1) Odontophorinae, hypotarsus usually 
with two roofed calcaneal canals; Phasianinae, hypotarsus with 
one roofed calcaneal canal. (2) Odontophorinae, inner calcaneal 
ridge without distal extension; Phasianinae, inner calcaneal ridge 
usually with distal extension that runs two-thirds the way down 
shaft. (3) Odontophorinae, posterior shaft usually without spur 
core; Phasianinae, posterior shaft usually with spur core, or rudi- 
mentary spur core in male. 

It is interesting to note that Sibley (1960) remarked that on the 
basis of electrophoretic patterns of egg-white proteins the New 
World quails might represent a separate subfamily. 

Summary 

On the basis of postcranial osteology the Cracidae (curassows), 
Megapodiidae (mound-builders), and Opisthocomidae (hoatzins) 
are much more closely related to each other than to the remaining 
groups of the Galliformes. These three families are considered to 
be primitive based on the thesis that the Cracidae comprise the 



Holman: Osteology of Galliformes 251 

most primitive gallinaceous family. The Opisthocomidae show 
many skeletal specializations. 

Among the remaining groups, the families Numididae (guinea- 
fowls), Meleagrididae (turkeys), and Phasianidae (with subfamilies 
Tetraoninae, grouse; Phasianinae, pheasants, junglefowls, peacocks, 
Old World quails; and Odontophorinae, New World quails) are 
recognized. 

The Numididae and Meleagrididae retain a few primitive skel- 
etal characters. In general, the Phasianidae are more advanced. 
The New World qauils (Odontophorinae) are considered to be the 
phasianid group that has departed most radically from the primi- 
tive cracid-like condition. 

Literature Cited 

Ashley, James F. 1941. A study of the structure of the humerus in the 
Corvidae. Condor, vol. 43, no. 4, pp. 184-195, 7 figs. 

Brodkorb, Pierce. 1964. Catalogue of fossil birds: part 2 (Anseriformes 
through Galliformes). Bull. Florida State Mus,, vol. 8, no. 3, pp. 195- 
335. 

Holman, J. A. 1961. Osteology of living and fossil New World quails 
(Aves, Galliformes). Bull. Florida State Mus., vol. 6, no. 2, pp. 131- 
233, 3 figs. 

Howard, Hildegarde. 1929. The avifauna of the Emeryville shellmound. 
Univ. Calif. Publ. Zool., vol. 32, no. 2, pp. 301-394, 4 pis., 55 figs. 

Hudson, G. E., P. J. Lanzillotti, and G. D. Edwards. 1959. Muscles of 
the pelvic limb in galliform birds. Amer. Midland Naturalist, vol. 61, 
no. 1, pp. 1-67, 20 figs. 

Mainardi, Danilo. 1960. Immunological relationships of the peacock (Pavo 
cristatus). Misc. Rept. Yamashina's Inst. Ornith. Zool., vol. 35, pp. 
130-132. 

. 1963. Immunological distances and phylogenetic relationships in 

birds. Proc. XIII Internat. Ornith. Congr., pp. 103-114. 

Mainardi, Danilo, and F. Guerra. 1959. Anticorpi naturali di siero bovino 
contro antigeni eritrocitari di Galliformi. Boll. Soc. Ital. Biol. Sperim., 
vol. 35, pp. 1805-1806. 

Mainardi, Danilo, and A. M. Taibel. 1962a. Studio immunogenetico sulle 
parentele filogenetiche nell'ordine dei Galliformi. Rendiconti Istituto 
Lombardo, vol. 96, pp. 131-140. 

. 1962b. Filogenesi dei Galliformi. Op. cit., vol. 96, pp. 118-130. 



252 Quarterly Journal of the Florida Academy of Sciences 

Mayr, Ernst, and D. Amadon. 1951. A classification of Recent birds. 
Amer. Mus. Novitates, no. 1496, pp. 1-42. 

Ridgway, Robert, and Herbert Friedmann. 1946. The birds of North 
and Middle America. Bull. U. S. Nat. Mus., no. 50, pt. 10, pp. 1-484, 
28 figs. 

Sibley, Charles G. 1960. The electrophoretic patterns of avian egg-white 
proteins as taxonomic characters. Ibis, vol. 102, no. 2, pp. 215-259, 
15 figs. 

Stresemann, Erwin. 1959. The status of avian systematics and its un- 
solved problems. Auk, vol. 76, pp. 269-280. 

Taibel, A. M. 1961. Analogie fisio-etologiche nel settore riproduttivo tra 
Afropavo Chapin e Penelope Merrem. Riv. Sc. Nat. 'Natura' vol. 52, 
pp. 57-64. 

Tordoff, Harrison B., and J. B. Macdonald. 1957. A new bird (Family 
Cracidae) from the early Oligocene of South Dakota. Auk, vol. 74, no. 
2, pp. 174-184, 1 fig. 

Wetmore, Alexander. 1960. A classification for the birds of the world. 
Smithsonian Misc. Coll., vol. 139, no. 11, pp. 1-37. 

Department of Biological Sciences, Illinois State University, 
Normal, Illinois. 

Quart. Jour. Florida Acad. Sci. 27(3) 1964 



FLORIDA ACADEMY of SCIENCES 

Institutional Members 
for 1964 

Barry College 

Florida Presbyterian College 

Florida Southern University 

Florida State University 

General Telephone Company of Florida 

Jacksonville University 

Miami-Dade Junior College 

Mound Park Hospital Foundation 

Rollins College 

Stetson University 

University of Florida 

University of Miami 

University of South Florida 

University of Tampa 



The next annual meeting will be at Tallahassee, March 11-13, 
1965. The chairman of the committee on local arrangements is 
Dr. Ruth S. Breen, Department of Botany, Florida State University. 



FLORIDA ACADEMY of SCIENCES 

Founded 1936 



OFFICERS FOR 1964 

President: George K. Reid 

Department of Biology, Florida Presbyterian College 

St. Petersburg, Florida 

President Elect: O. E. Frye, Jr. 

Game and Fresh Water Fish Commission 

Tallahassee, Florida 

Secretary: John D. Kilby 

Department of Biology, University of Florida 

Gainesville, Florida 

Treasurer: John S. Ross 

Department of Physics, Rollins College 

Winter Park, Florida 

Editor: Pierce Brodkorb 

Department of Biology, University of Florida 

Gainesville, Florida 



Membership applications, subscriptions, renewals, change 
of address, and orders for back numbers should 
be addressed to the Treasurer 



Correspondence regarding exchanges 
should be addressed to 

Gift and Exchange Section, University of Florida Libraries 
Gainesville, Florida 






Quarterly Journal 

of the 

Florida Academy of Sciences 



Vol. 27 December, 1964 No. 4 

CONTENTS 

Stability of ketones toward allcyl borates Carl H. Snyder 253 

An entropy function Vernon E. Derr 255 

A new barnacle from the Tamiami Miocene Arnold Ross 271 

Type locality of Platylepas tvilsoni Ross Arnold Ross 278 

Glucose nutrition and longevity in oysters 

Larry Gillespie, R. M. Ingle, and Walter K. Havens 279 

Notes on postlarvae of Panulirus argus 

Roy Witham, Robert M. Ingle, and Harold W. Sims, Jr. 289 

Two dragonflies new to Florida Dennis R. Paulson 298 

New Atlantic coast ranges for fishes 

William D. Anderson, Jr., and Elmer J. Gutherz 299 

Hypertensive effect of Latrodectus venoms 

John D. McCrone and Roger J. Porter 307 

A new glass lizard from Veracruz, Mexico /. Alan Holman 311 

Subspeciation in Sphaerodactylus copei 

Albert Schwartz and Richard Thomas 316 

Isolating mechanisms in snakes Wilfred T. Neill 333 

Large quahog clams from Boca Ciega Bay Harold W. Sims, Jr. 348 



Mailed February 26, 1965 



Quarterly Journal of the Florida Academy of Sciences 
Editor: Pierce Brodkorb 



The Quarterly Journal welcomes original articles containing 
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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY OF SCIENCES 



Vol. 27 December, 1964 No. 4 

STABILITY OF KETONES TOWARD ALKYL BORATES 

Carl H. Snyder 

The reported lack of reactivity of alkyl borates in the Meerwein- 
Ponndorf-Verley (MPV) reduction of ketones (Kuivila et al., 1951) 
has been attributed to a highly effective back-coordination of the 
filled 2p orbitals of the oxygen atoms with the empty 2p orbital of 
the boron, thus effectively decreasing the coordinating power of 
the boron atom (Jackman and Macbeth, 1952). Although aluminum 
is an inherently weaker Lewis acid than boron, aluminum alkoxides 
are quite reactive in the MPV reduction. Because the empty 
p orbital of the 

3RCR + AKDCHfg, ^-> 3RCHR + 3RCR + Ai(0H) 3 

aluminum atom is at the third quantum level, back-coordination 
with the oxygen 2p electrons is ineffective in these compounds and 
the empty aluminum 3p orbital is available for participation in the 
reaction. 

Since a similar disparity of quantum levels exists in the boron- 
sulfur bond, reaction of thioborate esters in the MPV reduction 
might be anticipated. Mikhailov and Fedotov (1961) report that 
thioborate esters do react with ketones, but to give thioketals and 
boric oxide rather than alcohols and thioketones. 

9 , R's, ,SR' 

3RCR + 2B(SR) 3 > 3RCR + B z 3 

Kuivila's reaction diagnostic was distillation of low molecular 
weight, highly volatile aldehydes or ketones produced in the re- 



msftotion MAR 5 1965 



254 Quarterly Journal of the Florida Academy of Sciences 

duction. Thus reported absence of a highly volatile product would 
not differentiate between unreactivity of the reagents and reactivity 
to produce relatively high molecular weight ketals and boric oxide 
by a process similar to that reported by Mikhailov and Fedotov. 
Accordingly a portion of Kuivila's work was repeated, but with 
direct analysis of reaction mixtures for unreacted ketones. Reflux- 
ing 0.05 mole of 2-butanone, cyclopentanone, or cyclohexanone 
with 0.1 mole of methyl, ethyl, or isopropyl borate for 5 hr. under 
nitrogen followed by dilution with heptane, gas chromatography 
(20 per cent Carbowax 20M on Chromosorb P) and comparison with 
standard solutions led in all cases to 91-98 per cent recovery of 
starting ketone. There was no evidence of ketal formation. Thus 
stability of aliphatic ketones toward alkyl borates is confirmed and 
the possibility of ketal formation is eliminated. 

Literature Cited 

Jackman, L. M., and A. K. Macbeth. 1952. Reductions with aluminum 
alkoxides. Part III. The kinetics of the racemization of optically ac- 
tive alkoxides by their corresponding ketones. Jour. Chem. Soc, pp. 
3252-3260. 

KurviLA, H. G., S. C. Slack, and P. K. Siiteri. 1951. Reactions of alde- 
hydes and ketones with alkyl borates. Jour. Amer. Chem. Soc, vol. 73, 
pp. 123-124. 

Mikhailov, B. M., and N. S. Fedotov. 1961. Reactions of esters of thio- 
boronic and organothioboronic acids with carbonyl compounds. Izv. 
Akad. Nauk S. S. S. R., Otd. Khim. Nauk., pp. 999-1001; Chem. Abst, 
vol. 57, p. 16643i. 

Department of Chemistry, University of Miami, Coral Gables 
46, Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



AN ENTROPY FUNCTION 
Vernon E. Derr 

Entropy, or rather its negative, was introduced into statistical 
mechanics by Boltzmann in order to show that any non-equilibrium 
situation would tend toward equilibrium. A discussion of the 
history of various proposed entropy functions and a critical anal- 
ysis are given by ter Haar (1955) and will not be reviewed here. 
However, it is desirable to point out that most suggested quantum 
mechanical entropy functions suffer from one or more defects, such 
as i) not changing with time, or ii) the entropies of two or more 
isolated systems are not additive, or iii) they have no physical con- 
tent but are the direct result of an application of Klein's lemma 
(Tolman, 1950). 

Speaking more positively we ask that an entropy function 
should: 

1) indicate by a change in time, the approach of a system to- 
ward equilibrium, tending to a fixed number at t— »oo; 

2) be additive, i.e., the sum of the entropies of two non-inter- 
acting systems should be equal to the entropy calculated by 

i considering the systems as a single system (we will see that 
it is necessary to add that the entropy to be presented will 
only be additive when the systems are in equilibrium); 

3) be dependent on the number of states in the system. 

It will be seen that the entropy function to be defined will fulfill 
these requirements. 

We begin by recalling that p(t) = U p(o) U" 1 = T p(o), where 

[ex. CX^~\ 

C C ) which 

is the ensemble average of the pairs of products of the quantum 
mechanical probability amplitudes for the states labeled by the 
subscripts. We see that if T is dependent on a single parameter 
(t = time) then we may write P ((t) = T^^o). Then, if T(t x + 
t 2 ) = T(t 2 )T(ti), p(ti + t 2 ) = T(t 2 )f (ti) P (o). If T depends only on 
the time interval and not on the setting of t = 0, then T(t 2 )T(ti) = 
T(t 2 + ti), and if t ± = t 2 = t, T 2 (t) = T(2t). The operator T will 



256 Quarterly Journal of the Florida Academy of Scdsnces 

b b 

depend on the interval only, if and only if / A(t)dt [where / 

a a 

A^dt 1 = U(t)] is a function of b — a, and not of a, b, separately. 
This will occur in case the system is isolated, and can occur under 
special conditions on A(t) and b — a, for example, periodicity of 
A(t). At first we will assume that T depends only on the time in- 
terval, later discussing the case where 8, the time derivative of the 
external parameter, is not zero. We see now that p(t) = Tp(o), 
p(2t) = T 2 p(o). . . etc., and we may form the time average 

T m p + T m+ V + lp + ... + T n - 1 p 

mn n - m q\ 

where p is the density matrix at some time t m and T is the operator 
which gives the density matrix at a later time t, i.e., p(t m +i) = 
T (t)p(t m ), where t = t mn — t in . We will frequently let m = o. 
Thus <f> mn is a time average of the density matrices which describe 
the system at the time t m , t m +i, . . . , t n -i. 

It follows immediately from the definitions of T and p that Tp 

. / P + Tp _ 
is a density matrix. Further, the tn 2 •' " ' (where tr is 

the trace) since 



tr(^-^) = 1/2 trip + Tp) 



= 1/2 [tr(p) + tr(Tp)] = 1. 

We know (Riesz and Nagy, 1953) that <£ nm tends to a limit, call 
it 6, as (n - m)— > 00 . From the considerations of completeness we 
know that belongs to the Hilbert space of matrices of bounded 
norm and that TO ±= 0, i.e., 6 is invariant under the operator T. 
But T0 =" U0U- 1 — so that U0 = ^U; that is, commutes 

, -27TJHt 

with U. But { h } (Tolman, 1950, Chap. 5, Sec. 96), 

U = e 

where H is the time independent Hamiltonian of the system. 
Hence, since if 6 commutes with U, it commutes with H, we see 
that the limit of the average cf> nm as (n - m)-> 00 , commutes with H 
and is a density matrix which is a stationary solution of the Liou- 



Derr: An Entropy Function 



257 



ville equation ££= __JL(pH - Hp). For we see that if p and 

H commute, -^r- = 0. Thus the class of density matrices which 

are limits of our time average is identical with the class which are 
stationary solutions of the quantum mechanical Liouville equa- 
tion. 

One might hope that more would be true, for example that 
if the limit of <j> nm exists, then it might be equal to the asymptotic 
solution of the Liouville equation. However, there does not seem 
to be any result of this kind possible. Hence, the Liouville equa- 
tion is not useful in describing the approach to equilibrium. 

Let us consider more carefully the limit 0. As is shown by 
Halmos (1956), the limit 6 is the projection of on the manifold 
of elements of Hilbert space which are invariant under T. i.e., 6 
belongs to the set of (f ePf = f) where P is the projection opera- 
tor. We may gain an intuitive idea of the content of this theorem 
by considering a three dimensional case. Let T be the operator 
which rotates a vector about the Z axis. Then the invariant sub- 



,ii 



7 



\ 



\ 



/ 



\ 



/ 



o 



/ 



_v 



/ 



X 



258 Quarterly Journal of the Florida Academy of Sciences 

space of the rotation is the Z axis, and the space orthogonal to 
it is the x, y plane. A vector f may be represented as a sum of a 
vector (I) from the invariant subspace and a vector (Q) from the 
x, y plane. Now let the rotation occur. The vector I remains un- 
changed, while Q rotates in the x, y plane. The rotation results 
in a new vector in the x, y plane, since we assume T is not the unit 
operator. Whether or not T j Q = Q for some j, the average of the 
vectors obtained in the x, y plane tends to zero. Thus the average 
of the f vector and its rotations is equal to I which is the projec- 
tion of f on to the invariant subspace, the Z-axis. In the case of 
the higher dimensionality of a Hilbert space, the picture is similar. 
It is clear from this picture that does not approach the invariant 
axis, but rotates around it. 

As shown by Hopf (1948) the limit 6 is given by the eigen- 
function of T corresponding to the zero eigenfrequency, i.e., 

s = A where Td> = e <b for X = 0, when X = is a simple 

eigenfrequency, that is, there is no degeneracy. In the case where 
T<f> is U<^U _1 , it is clear that degeneracy always occurs because in 

the case of A = 0, we have iH , -iH 

e 4>e =9 

or 

iH x ^ iH 

and <f> will commute with e iH if and only if <f> is any function of H. 
Therefore, we have the result that the class on invariants to T 
is the set of matrices f(H), where we restrict these to be analytic 
functions. Further it is clear that not all matrices f(H) are per- 
missible, since the limit matrix must have trace 1, and its di- 
agonal elements must be non-negative. Since = P p , we have that 

6 = cf(H) 
where c is a constant (scalar) chosen so that Xxd = 1. It is clear 
from these considerations that no one function from the set f(H) 
is favored as the limit function. The microcanonical distribution 

-H 
■K- e > K a constant parameter, is a possible candidate for 6, but 

there is no necessity that it will occur. Insufficient information is 

given to decide what the limit will be, even if the density matrix 

P and the operator T are given, since T is in general degenerate 

for the eigenfrequency A = O. 



Derr: An Entropy Function 259 

We have now completed the discussion of the essential prop- 
erties of the time average um which will be needed for the discus- 
sion of the entropy function. We define the entropy function 

a - tr(d> i n 6 ) 



or 



t^pCp) 



« N = 4 p(o)+Tp(o) ;- + 



+T N- 

tn P(o) p(o)+,..+ T i N 



This is a fine-grained function, that is it makes use of the fine- 
grained probabilities rather than any coarse-grained probabilities. 
We proceed to develop its properties. 

Properties of <t n 

The limit, as t-»oo, i.e., N-*<x of <r N , exists. This is evident since 

lim 4> exists arid is not zero. Recall that lim <J> is a density 
N-*oo N—oo 

matrix. We may diagonalize simultaneously 8 _ lim <|> and In 8, 

i.J > ;« , " N-00 N 

leaving the tr(0 In 8) unchanged, i.e., tr(U0U Jl (U In 8 U 1 ) = 
tr(0 In 8). Let D = U 8 V Jl be diagonal. This is always possible 
since 8 is Hermitian. The entries on the diagonal (Da) are the 
(positive) eigenvalues of 8, arid % Ki == 1. Thus tr(D In D) == 
SiDiilnDii. 

If Si and S 2 are two given, isolated non-interacting systems, 
with density matrices p w and p (2 \ and 



a) t / VD + Tp <i> + ... + T N-yi> 

a N =tr l N 



. P (1) + - + T 

in 



N 



N-l (1)\ 

— ) 



260 Quarterly Journal of the Florida Academy of Sciences 
and . ... „_ x 2 

P 



(2) t / p< 2 » t .,. + T S - 1 
f N = H N 



P (2) +...+ 

N 



are the entropies respectively at a given time of the two systems, 
then the entropy of the combined system S = Si plus S 2 is 



a N = tr 



p xp . +T(p xp )+...+ T (p xp ) 



N 



p xp +...+ T (p xp ) | 
N J 



where p (1) x p (2> is the direct product of the density matrices of the 
separate systems. 

Here we start with a simple example in order to define the 
direct product. Consider that the density matrices for the systems 
are diagonal, 

and let 



(1) 



O 



ind p 



(2) 



O 



The a's and b's are the probabilities of the states of the systems 
1 and 2 respectively. The systems are assumed non-interacting 
and statistically independent, hence the probability of a combined 
state, i.e., for example the probability that Si is in state i and S-. 
is in state j, is aibj. Thus, the density matrix describing the com- 
bined system Si is: ; 



O 



a 2 b 2 



a l b 2 



a 2 b l 



Derr: An Entropy Function 261 

where the entries on the diagonal are every possible product of 
pairs of the a's and b's. For diagonal matrices the matrix p is 
called the direct product of the matrices p w x p (2) , = p (1) x p <2 \ 
Consider <y 1 = tr(p \np), where p is the (diagonal) density matrix 
for the combined systems, S = Si plus S 2 . 



Then 



f / (1) (2) „ (1) (2), 
<J 1 = trip xp inp X p ) 

= 2. . a.bjna.b. 
M i] i 3 

= 2. . a.b. in a. + 2. . a.b. in b. 
i.J i 3 i 1.3 i 3 3 

= 2. a. in a. (2b.) + 2.b. in b (2. a ) 
11 13 3 3 3 l i 

Hence using the fact that 2. b. = 2. a. = 1, we see 

2 2 i 1 

^-■tHp^inp^+W^inp^)'. 

or 

Vr - n- (1) 4- (2) 

^-^l +CJ 1 * 

Thus the entropy at t = of the combined system is equal to 
the sum of the entropies of the separate subsystem. 

Our next task is to see if this result can be extended to ; o-^ 
and to non-diagonal p's. We first hold N = 1 as above and investi- 
gate the relation between '■■■■''■ 

, (1) A (2)> (1) A (2) 

a and a and cr for p and p 

non-diagonal matrices. Then 

<j = tr(pinp) = tr [p (1) x p (2) inp (1) x p (2 ^ , 

and since we can diagonalize without changing the value of the 
trace 



262 Quarterly Journal of the Florida Academy of Sciences 

V .r(D (1 » X D ,2) ,„D (1) xD |2) , 

= tr(D (1) ,„D (1) ) + tr(D (2) in D (2) ) 

-„ (1) + „ < 2) 
" ff l + CT 1 

So in this case also the entropy of the combined system is the sum 
of the entropies of the separate subsystems. 

Now consider the general case. Let P w and p (2) be the density 
matrices for two non-interacting systems. Let T (1) and T (2) be the 
unitary operators on P a) and p (2) respectively which govern the 
evolution of the density matrix in time. Let p = P a) X /° <2> be the 
density matrix of the systems, considered as a single system. We 

m (1) 1 

have that x p(o) = U p(o) U = p(t), an d similarly for the 

second system. Further, Tp is (Ui X U 2 ) p (Ur 1 X U 2 _1 ), (Murn- 
aghan, 1954). We define the entropy as a function of time by: 

cr (l) = tr [(4> (l) in c|> l )]and a = tr[(4>in <j>)J 

where 

_ p (i) + T (i) p (i) + ... + T (i)N-l p (i) 

*i N 

We ask, under what conditions does o- = o- (1) + o- <2) ? 

In order to apply the proof used previously, it is necessary to 
transform the matrices <f> into diagonal matrices in such a way that 
<r is unchanged. This, of course, may be accomplished by a unitary 
transformation which leaves the trace unchanged. However, more 
is needed, since if a = tr(DlnD), D a diagonal matrix, in order to 
insure additivity we must have that D = D (1) x D (2) , where D (1> 
is the matrix obtained by diagonalizing <£ (i) . This in general may 
only occur when we may simultaneously diagonalize p <l) , T (l> 
p (i) , etc. Thus we assume that they may be simultaneously diag- 
onalized, leaving till later the discussion of the significance of this 
assumption. 

It is clear that if p w is diagonalized in the form S (1) p (1) S (1)_1 



Derr: An Entropy Function 263 

and similarly for p i2 \ then p w X <2> * s diagonalized in the form 
(S (1) X S <2) ) ( P w X p (3) ) (S (1)_1 X S^" 1 ). We will assume that a 
sufficient condition that two infinite Hermitian matrices are simul- 
taneously diagonalizable is that they commute. Thus we now 
apply a unitary transformation to <f> which diagonalizes simulta- 
neously p, Tp . . . etc ., i.e., 

a = tr(<|)in<t>) = tHS^S"" 1 in S^S -1 ) 



or 



. J SpS" 1 
a = tr — 



+ STpS" 1 +...+ ST N_1 pS" 1 



in 



N 
SpS" 1 



1 + STpS" 1 +...+ ST 1 *" 1 pS" 1 

N J' 



We remark that p and Tp (and hence x^p > N = 2; 3 — ) have 
the same eigenvalues, when T is unitary, and thus 

. rlL(l) q (2) w (l) (2) u _(l)-l _(2)-l 
<j> = N [(S x S ,)(p x p )(S x S 



+ ... + (S (1) xS (2) )(T (1) p (1) xT (2) p (2) ) 



-ll 



(S VA/ xS w x ) 



1 D (1) xD (2) +... + D (1) xD (2) n (l) (2) 

<*>=• Jj =D xD . 



But 

■ (i) _ s (1) p< 1 V 1 >- 1 ,...,s (1 ¥ 1) p (1) s (1) - 1 

N 



Hence 

(1). (2) 

ct = ct + cr , 



264 Quarterly Journal of the Florida Academy of Sciences 



since 



a = tr(D (1) xD (2) i„D (1) xD (2) ) 
= tr(D (1) inD (1 >) + tr(D (2 >i„D (2) ), 



and this is the result sought. 

What is the significance of the assumptions that p and Tp com- 
mute? That they commute means 

p(Tp) = (Tp)p 



or 



or 



pUplT 1 = Up U _1 p 



iHt -iHt iHt -iHt 
p e pe =_e pe p. 

This implies an especially simple relationship between p and Tp; 
namely, that one is a function of the other. In the event that p is 
diagonal, Tp must also be diagonal. As a special case is included 
that where p is stationary under the T operation, i.e., the system 
is in equilibrium. 

Thus although the quantity a, defined to be the entropy, is not 
always additive, it is so when the systems are in equilibrium. 

Let p (o) be an arbitrary density matrix. Let U be diagonal 
and unitary. We will compute ii m CT for this case. First we 

N 
N—oo 

see that if U is diagonal and unitary, 



u n 



U 22 



U 



U 23 



O 




Derr: An Entropy Function 265 



then u 11 v 11 = 1, and u jj = v xl> hence U is of 
form 



e^l °\ /e~ i<p l 

U= i4> 2 \andU" 1 



\0 
Then Up(o) U~l is: 




^(o) e i( *l"^) pi2(o) 



e i(+2- + i)P2i(o) p (o) 



Thus 

p,(o)+e +i ^e"^p : Jo) 



[/ \ _l tt / xtt- 1 ! P..(o) + e Y i e \j p. J 
p(o) + Up(o)U r ij ;_ij_ 

2 J .. " 2 



ij \ 2 

P kk (o) if i = j = k 



= V o) t 2 ]■**/ 

We may similarly compute: 

f TT N , , TT -Nl , . iN(4>,-<t>-) 

[U p(o)U J .. = p..(o) e Tl T J . 



i] 1.1 



266 Quarterly Journal of the Florida Academy of Scrences 
Thus: 

rdl i _ rp(o)-mp(o)u~ 1 +...+ u N " 1 P (o)u- (N " 1 > | 

1>J " L n— J. 



= p 

and obviously 



(o) ri + e i( V^> + ,..fe i(N - 1 ^i-^) | 



f p kk (o) lf 1 = 3 = k) 
4> =lim(N-oo) py = 1 tt I 

L -J ij I if i ^ j. ) 

if we assume no degeneracies, i.e., 4>i ^ <j>. if 
i ^ j. Thus J 



lim or = cr = tr(4>in<|>) = S (p (o) in p. . (o)). 

N-*oo k kk kk 



We know from the theorem previously quoted that the limit <£ 
always exists. Hence we have here an entropy function which 
starts at a value tr [ P (o) In p (o)] and tends to a value 2 k [pkk(o) In 
/okk(o)]. The off-diagonal terms, which contain the phases of the 
states, although not initially zero, lose their effect on the entropy 
as time tends to infinity. Note that the initial entropy tr[p (o) In p 
(o)] can be written as 2 k [A k In A k ] where the A's are the eigen- 
values of the density matrix p(o). Speaking broadly, the value of 
o- will differ from that of tr[p (o) In p (o)] if the eigenvalues of p(o) 
are "spread" more than the diagonal terms. We will attempt to 
consider this question more carefully. 

We will compare the values of o-i = tr[p (o) In p (o)] and 2k [?kk 
(o) In /okk(o)] = o-oo, as above. Recall that a 1 = 2k A k In A k where 
the A's are the eigenvalues of the density matrix P (o). Let us first 
consider the special case where 



Derr: An Entropy Function 



267 



p(o) = 



P n P 12 o 



P 21 P 22 ° 



° P 33 ° 



44 



that is, of the off-diagonal terms, only p 12 and p 2 i are not zero. In 
this case we may obtain the eigenvalues of p (0) easily. In addi- 
tion to pss, p44, . . . the other eigenvalues are the solutions of: 



(P U- X) "l2 



(P 21 ) (P 22 " X) 



from which 



= -0, 



X ± 



P ll +P 22 



± V 1/4(P 11^22 )2+ K 



2 f -""11 r "22' ^12 

The difference of these eigenvalues is 

2 



+ 



yi/4(p n - Poo ) 2 + ( Pio ) 2 



22 ; 



12' 



V (P ll- p 22 )2+4<p 12 )2 



Thus, the difference of these eigenvalues is greater than the differ- 
ence of pn and p 22 - Let us compare for this case o-i and cr . 
Since all the eigenvalues except the first two are the same 



as P kk (o), k = 3, 4, ..., we have 



268 Quarterly Journal of the Florida Academy of Sciences 
o-oo-o-l =^ + inX ++ A_inX_-p ii inp ll -p 22 inp 22 . 
But 

X + + \_ = P :1 + P 22 = k, say, and 

(\-^)>(p u -p 22 ). 

Under these conditions, the concave function 

gives us that a < cr-. . 
oo x 

The question remains open however as to whether this result 
can be extended to arbitrary P (o). We do know that if the prob- 
abilities of the states, Pkk , are zero for k > N initially then there 

exists a class of density matrices for which ^ < °y This can be 

seen by letting the off diagonal terms be almost zero except for 
the terms p 12 and p 2 i. Then, by continuity, the eigenvalues of 
the matrix are almost 

N 
V X -'<>33' p 44'" ' and k S = /kk n \k 

N 
can be made to differ only slightly from the 2 X n X whereas, 

k=2 k k 

since p 12 2 is not small, we may, as before, see that cr^ <( ?y 

Thus we see that there is a class of density matrices for which 

oo l' and thus the entropy in these cases changes with 

time. For the system just examined, it does not necessarily go to 
a minimum, for there can be no changes in the probabilities of the 
states since our operator T was diagonal. 

We now consider that a random perturbing force acts on the 
system through an external parameter a. We assume the perturba- 
tion is very small and that the energy it may impart to the system 
is bounded, i.e., tr(Hp) where H is the unperturbed Hamiltonian, 
does not grow beyond all bounds. The perturbation is considered 



Dekr: An Entropy Function 269 

to be so small that the identity of the energy levels of the system 
is retained, i.e., E n (a) is almost equal to E n (a ) where the latter is 
the energy level for zero perturbation. More precisely E n (a) — 
E n (a ) << E n (a ) ~ E„ ± ] (a ). We wish to consider the time 
course of the entropy o-. 

If the random perturbing force is not present and we consider 
that the density matrix is in an energy representation, then the 
diagonal terms do not change, but the off-diagonal terms in 

i(4> -4> ) 
general are complex numbers of the form r r e n m 

i(E -E )t n m 

e n m and are rotating vectors in the complex plane. The 
effect of the operator T where it contains the perturbation is to 
change the lengths of the vectors which are the off -diagonal terms, 
as well as the phases, and to change the diagonal terms. The 
effect on the off-diagonal terms will be to replace a set of vectors 
of equal length but random orientation by a set having random 
length and random orientation. When the time average </> N is 
considered we see that the effect will be to make the off-diagonal 
terms of the limit matrix 6 be zero. The diagonal terms of course 
do not all tend to zero being the average of non-negative real 
members, whose sum is one. 

It does not seem possible here as well as in the previously 
considered deterministic case to find the precise form of the limit 
matrix 0, because in addition to the degeneracy problem, there is 
additional uncertainty due to the random perturbation. However, 
if we make the assumption that the effect of this perturbation, on 
the average, is to reduce any locally maximum state probabilities, 
we see that the limit density matrix 6 will have the following 
properties: 

a) Si pa = 1, pa > 0. 

b) /oh — » as i -» oo sufficiently fast so that S n = i n pnn con- 
verges, c) No states have zero probability, i.e., pa not = 0, i = 
1,2, .. . d) Our assumption that there are no local maxima, re- 
quires that pii as a function of i be non-increasing. 

Thus the general character of the diagonal terms of the limit 

matrix 6 is similar to the canonical distribution, j^e" . 



270 Quarterly Journal of the Florida Academy of Sciences 

In conclusion we may observe that although previously de- 
fined entropy functions have some of the characteristics listed in 
the introduction, no one of them possesses all of the properties 
necessary for a function to describe the passage of a system to 
equilibrium. The function o- defined herein is additive, dependent 
on the number of states, and measures the change with time of 
a system tending toward equilibrium. Further, although exact 

-E. 

equality to the canonical distribution Ke l is not necessarily as- 
sured in equilibrium, nevertheless the energy dependence is given 
by a class of non-increasing functions similar to the canonical dis- 
tribution. Finally the function o- depends on the fine-grained prob- 
abilities of the quantum mechanical states and thus contains all 
the details that a maximal observation of the system could achieve, 
in an averaged form. 

Literature Cited 

Halmos, Paul R. 1956. Lectures on ergodic theory. Mathematical Society 
of Japan, Tokyo, 99 pp. 

Hopf, Eberhard. 1948. Ergodentheorie. Chelsea Publishing Co., New 
York, 83 pp. 

Murnaghan, Francis D. 1938. The theory of group representations. Johns 
Hopkins Press, Baltimore, 369 pp. 

Riesz, Frederic, and Sz.-Nacy, Bela. 1953. Lecons d'analyse fonction- 
nelle. Academie de Sciences de Hongrie, 445 pp. 

ter Haar, D. 1955. Foundations of statistical mechanics. Rev. Modern 
Physics, vol. 27, no. 3, pp. 289-338. 

Tolman, Richard C. 1950. Principles of statistical mechanics. Oxford 
University Press, London, 661 pp. 

Martin Company, Orlando, Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



A NEW BARNACLE FROM THE TAMIAMI MIOCENE 
Arnold Ross 

The present study is the fourth in a series on the fossil balano- 
morph Cirripedia of Florida. Herein the author describes a new 
species of Balanus from the Tamiami Formation, of Late Miocene 
age, in southwestern Florida. 

Based on Florida State Museum collections at the University of 
Florida the author has thus far been able to identify the following 
Tamiami species: Balanus trigonus Darwin, B. glyptopoma Pilsbry, 
B. concavus proteus Conrad, Chelonibia testudinaria (Linnaeus), 
and C. patula (Ranzani). 

The stratigraphic and zoogeographic distribution of Chelonibia 
was presented by Ross (1963). Balanus trigonus has not heretofore 
been reported from the Miocene of the southeastern United States. 
The two specimens of supposed Late Miocene age from the York- 
town Formation in Virginia (Ross, 1964) are more probably Pleisto- 
cene, if not Recent. Balanus glyptopoma ranges from Florida to 
Virginia (Pilsbry, 1916), and is here believed to be restricted to 
Miocene sediments. Specimens referred to this species by Pils- 
bry (1918) from the Panama Canal Zone probably represent an 
undescribed species. Balanus concavus proteus ranges from Mary- 
land to Florida. Specimens of this species described by Cushman 
(1904) from Gay Head, Massachusetts, must be re-examined before 
drawing any conclusions regarding their taxonomic assignment. 

Cornwall (1962), in one of his numerous studies on interlaminate 
shell figures, reported the megabalanid Balanus tintinnabulum (Lin- 
naeus) from Miocene sediments in DeSoto County, Florida, pre- 
sumably from one of the Tamiami lithofacies. This record, how- 
ever, is not in accord with the author's findings based on morpho- 
logical studies over a period of five years; the figures presented by 
Cornwall (1956, 1959, 1962) were therefore restudied for some 
incongruity. Examination of the figure presented for B. tintinnab- 
ulum, PL 1, fig. 10 (1962), indicates a great similarity to those of PI. 
2, fig. 14 (1962), text-figs, lg, lh, 3i, 3j (1956), and PL 1, fig. 5 (1959), 
all of which are typical of the concavus complex. The pattern 
found in B. tintinnabulum and related forms, although similar, are 
distinct from B. concavus (sensu lato). Without adequate morpho- 
logical material no attempt should be made to base species identi- 



272 Quarterly Journal of the Florida Academy of Sciences 

fications wholly on interlaminate figures as Cornwall evidently 
did. 

Institutional abbreviations used herein are as follows: United 
States National Museum, U.S.N. M.; Florida State Museum, F.S.M. 

Family Balanidae Gray, 1825 
Subfamily Balaninae (Gray), 1825 

Genus Balanns DaCosta, 1778 
Balanus tamiamiensis, new species 

Diagnosis. The peritreme of the shell is strongly toothed and 
flaring. Transverse septa are present in the parietal and basal tubes. 
The peripheral lateral depressor muscle ridge of the scutum is com- 
posed of long, serrate teeth. 

Description. The shell is large, tubular, tubuloconic or conic, 
commonly flaring apically, and ornamented with high prominent 
ribs. There are, on the two paratypes, three ribs on the carina, one 
on the carinolateral, two on the lateral, and three on the rostrum. 
One of the paratypes (fig. li) has intercalated secondary ribs, but the 
number and position do not appear to be constant. The holotype 
(fig. la), on the other hand, appears to have been crowded and to 
have grown on an irregular substratum, and thus did not develop 
a normal rib pattern. The carinolateral compartment is approxi- 
mately one-third the width of the lateral compartment. The orifice 
is diamond-shaped to slightly pentagonal; the pentagonal form be- 
ing contingent upon the width and proximity of the rostrum to the 
laterals. The peritreme is very deeply toothed. The radii are 
moderate to very narrow, and bear both transverse and vertical stri- 
ations. They are also moderate to deeply sunken, and their sum- 
mits are both crenulated and steeply sloped. The lateral edges of 
the radii are septate, the septa denticulate along the basal portion 
only. The summits of the alae are sub-horizontal; those between 
the lateral plates and rostrum are more steeply sloped than those 
between the other compartments. The sheath occupies approxi- 
mately the upper one-half of the parietes. Its surface is orna- 
mented with strong vertical ridges, the surface of which bear im- 
bricating scales. The vertical ridges are most prominent on the lat- 
eral and carinolateral segments. The basal portion of the sheath 
depends freely, leaving a broad, deep pocket behind it. The in- 



Ross: New Miocene Barnacle 



273 




Fig. 1. Balanus tamiamiensis, n. sp. a, right lateral view of shell, holo- 
type, U.S.N.M. 648951 (actual height 46.4 mm), b, external view of left 
carinolateral, U.S.N.M. 648952 (actual height 45.1 mm), c, internal view of 
right lateral, U.S.N.M. 648953 (actual height 49.3 mm), d, external view of 
right scutum, holotype, U.S.N.M. 648951 (actual height 24.2 mm), e, inter- 
nal view of left scutum, holotype, U.S.N.M. 648951 (actual height 25.0 mm). 
/, right lateral view of shell, F.S.M. 1339 (actual height 35.9 mm), g, h, 
external and internal views of left tergum, holotype, U.S.N.M. 648951 (actual 
height 21.2 mm), i, apical view of shell, F.S.M. 1340 (actual width 34.8 mm). 



274 Quarterly Journal of the Florida Academy of Sciences 

ner surface of the shell is coarsely ribbed, the ribs extending upward 
into the hollow behind the sheath. The parietal tubes are large, 
rectangular at the base, and they are interrupted at irregular inter- 
vals by transverse septa. The radial tubes of the basis also bear 
transverse septa. At the periphery the basis is multicellular. 

The scutum is narrow and extremely high. The poorly pecti- 
nate occludent margin is approximately twice the length of the ter- 
gal and basal margins. Rather low, irregular, sinuous, imbricating 
growth ridges and poorly developed longitudinal striae ornament 
the external valve surface. The basi-tergal angle is evenly rounded. 
The inflected tergal segment is extremely narrow. The articular 
ridge is low and extends less than two-thirds the length of the ter- 
gal margin; the articular furrow is shallow and narrow. The ad- 
ductor ridge is moderately short, thin, undercut, and high, reach- 
ing its maximum above the middle. It terminates distally well 
above the basal margin of the valve. The depression for the in- 
sertion of the adductor muscle is deep, elongate-oval, and where 
immediately adjacent to the crest of the adductor ridge the periph- 
ery of the depression is strongly angular. A moderately high ridge 
occurs mediad and peripheral to the lateral depressor muscle de- 
pression. This depression extends apically about one-half the 
height of the valve. The marginal lip originates about one-third 
the distance above the basal margin and extends upward between 
the adductor and articular ridges. The crest of this ridge is not 
continuous, but rather broken into three or four distinct teeth. 
The pit for the rostral depressor muscle is narrow, elongate, and 
shallow. The apical third of the inner valve surface is marked by 
short, low, parallel to sub-parallel ridges. Similar, sparse ridges 
are found underneath the adductor ridge. Along the basal margin 
there are low, sub-obsolete vertical ridges which may prove to be 
due to corrosion. 

The tergum is narrow and high. Growth ridges on the exterior 
valve surface are not as prominent as those on the scuta. There 
appear to be sub-obsolete striae in the basiscutal region. The ex- 
ternal longitudinal furrow is closed on the apical one-half, open- 
ing gradually toward the basal portion. The growth ridges in the 
spur fasciole and longitudinal furrow are coarser than those on 
the valve proper. The carinal edge of the valve is inflected, the 
inflection being extremely narrow. The spur is moderately long, 
sub-angularly rounded distally, approximately one-fourth the width 



Ross: New Miocene Barnacle 275 

of the basal margin, and separated from the basiscutal angle by 
approximately its own width. The spur and scutad portion of the 
basal margin form an acute angle of about 70 degrees. The articu- 
lar ridge is low, but prominent. The articular furrow is broad, 
but shallow, and marked by a few, low, narrow ridges. There are 
six crests for the insertion of the tergal depressor muscle; they are 
low, inclined and moderately short. The inner surface of the valve 
is roughened with short, parallel ridges in the same manner as 
the scuta. 

Remarks. The general shape of the shell of Balanus tamiami- 
ensis somewhat approaches that of the Miocene to Recent east- 
ern Pacific species Balanus nubilus Darwin as figured by Pils- 
bry (1916, pi. 30, fig. la). The absence of transverse septa in the 
parietal tubes, complex sutural articulation surfaces, and the oper- 
cular characteristics of B. nubilus should serve to distinguish it 
from the present new species. 

Of the Atlantic Coastal Plain Miocene species in the concavus 
group this new taxon is closely allied to B. concavus proteus and 
B. glyptopoma. Balanus tamiamiensis may be distinguished from 
the former species by several morphological features, the most 
notable being the deeply toothed and flaring orifice, narrow and 
sunken radii, and transverse septa in the parietal and basal tubes. 
The scutum is much higher than either of these described species, 
and intermediate in width, B. glyptopoma being the narrowest. 
The external sculpture is finer and not nodose as in B. glyptopoma, 
but somewhat coarser than that of B. concavus proteus. The shell 
ribbing of B. glyptopoma is much finer than that of Balanus tami- 
amiensis. Although both species possess transverse parietal and 
basal septa, the narrower radii, flaring and deeply toothed orifice 
should aid in separating this new species from B. glyptopoma. 

Disposition of Types. The holotype shell (fig. la) and associ- 
ated opercular valves (figs. Id, e, g, h) are deposited in the collec- 
tions of the U. S. National Museum, catalogue number 648951. 
The two complete paratype shells, both lacking opercular valves, 
are deposited with the Florida State Museum, catalogue numbers 
1339 (fig. 1/) and 1340 (fig. It). The carinolateral and lateral com- 
partments, representing two specimens, are also deposited with the 
U. S. National Museum, catalogue numbers 648952 (fig. lb) and 
648953 (fig. lc). 



276 Quarterly Journal of the Florida Academy of Sciences 

Measurements of Holotype. Height of lateral compartment, 
46.4; carinorostral diameter, 44.2; carinorostral diameter of orifice, 
30.1; height of right scutum, 24.2; height of left tergum, 21.2 mm. 

Type Locality and Horizon. West Coast Rock Co. limerock 
quarry, S.W. Vi, Sec. 26, Township 45 S., Range 24 E., about 0.3 
miles west of U. S. Highway 41 on Florida Highway 865, and about 
8.0 miles south of Fort Myers, Lee County, Florida; "Buckingham" 
lithofacies, Tamiami Formation, Late Miocene; A. Ross collector, 
September, 1959. 

Habitat. Echols (1960), basing his observations on the Tamiami 
ectoproct bryozoans of the Sunniland area, Collier County, believed 
that the fauna indicated tropical or subtropical marine conditions. 
The echinoid (Kier, 1963) and molluscan (Mansfield, 1932) faunas 
also indicate warm hydrotemperatures for the Tamiami. 

Bathymetric conditions at this locality were postulated by Kier 
(1963), based on Arbacia crenulata, as being littoral. 

Etymology. The specific name tamiamiensis indicates that the 
species is from the Tamiami Formation of Florida. 

Acknowledgments 

The author is grateful to Dr. Harold K. Brooks, Curator of 
Invertebrate Paleontology, Florida State Museum, for making avail- 
able fossil material. He is indebted to Ronald J. Echols at whose 
home he stayed while in southern Florida and who accompanied 
the writer while collecting in this area. 

Literature Cited 

Cornwall, I. E. 1956. Identifying fossil and recent barnacles by the figures 
in the shell. Jour. Paleon., vol. 30, no. 3, pp. 646-651, text-figs. 1-3. 

. 1959. More shell figures and notes on barnacles. Canadian Jour. 

Zool., vol. 37, pp. 401-406, pis. 1-3. 

. 1962. The identification of barnacles, with further figures and notes. 

Ibid., vol. 40, pp. 621-629, pis. 1-2. 

Cushman, Joseph A. 1904. Miocene barnacles from Gay Head, Mass., with 
notes on Balanus proteus, Conrad. Amer. Geologist, vol. 34, pp. 293- 
296. 

Echols, Ronald J. 1960. The bryozoan fauna of the Tamiami Formation 
(Upper Miocene) of Florida. Unpubl. Masters Thesis, Univ. Florida, 
pp. 1-78, 1 fig. 



Ross: New Miocene Barnacle 277 

Kier, Porter M. 1963. Tertiary echinoids from the Caloosahatchee and 
Tamiami Formations of Florida. Smithsonian Misc. Coll., Publ. 4543, 
vol. 145, no. 5, pp. 1-63, text-figs. 1-58, pis. 1-18. 

Mansfield, Wendell C. 1932. Pliocene fossils from limestone in southern 
Florida. U. S. Geol. Surv., Prof. Paper 170-D, pp. 43-56, pis. 14-18. 

Pilsbry, Henry A. 1916. The sessile barnacles (Cirripedia) contained in the 
collections of the U. S. National Museum; including a monograph of 
the American Species. Bull. U. S. Nat. Mus., no. 93, pp. 1-366, figs. 
1-99, pis. 1-76. 

. 1918. Cirripedia from the Panama Canal Zone. Bull. U. S. Nat. 

Mus., no. 103, pp. 185-188, pi. 67. 

Ross, Arnold. 1963. Chelonibia in the Neogene of Florida. Quart. Jour. 
Florida Acad. Sci., vol. 26, no. 3, pp. 221-233, 2 figs. 

. 1964. Cirripedia from the Yorktown Formation (Miocene) of Vir- 
ginia. Jour. Paleon., vol. 38, no. 3, pp. 483-491, text-figs. 1-2, pis. 
71-72. 

Department of Geology, University of Florida, Gainesville, 
Florida. 



Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



TYPE LOCALITY OF PLATYLEPAS W1LSONI ROSS 

Arnold Ross 

In a letter dated October 25, 1963, Druid Wilson has called 
my attention to the incorrect citation of the type locality of Platy- 
lepas wilsoni Ross (1963), U. S. Geological Survey Cenozoic locality 
22805. The holotype was collected on the southwest side of Rim 
Ditch Canal, not the northeast side as stated. Furthermore, the 
type locality is approximately 175 yards northwest of the Florida 
East Coast Railroad bridge, not 500 yards northeast. The photo- 
graph of the type locality (op. cit, fig. 1) was taken approximately 
500 yards from the bridge. 

Literature Cited 

Ross, Arnold. 1963. A new Pleistocene Platylepas from Florida. Quart. 
Jour. Florida Acad. Sci., vol. 26, no. 2, pp. 150-158, figs. 1-3. 

Department of Geology, University of Florida, Gainesville, 
Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965): 278 



GLUCOSE NUTRITION AND LONGEVITY IN OYSTERS 
Larry Gillespie, R. M. Ingle, and Walter K. Havens 

The processes of growth and "fattening" are of fundamental im- 
portance to the oyster biologist and farmer. One of the objectives 
in oyster nutrition is to identify the various nutrients and feeds 
which can be utilized by this mollusk. The natural foods for the 
oyster are micro-organisms and perhaps detritus; however, the 
possibility that this shellfish is able to assimilate soluble substances 
such as carbohydrates should be considered. 

The objectives of this study were to investigate the utilization 
of glucose in the nutrition of the oyster. Also, observations were 
made on the effect of glucose as a stimulator to pumping and on 
the length of life of the oyster in artificial sea water. 

Review of Literature 

Yonge (1928) found that oysters were able to utilize glucose 
under abnormal conditions by indirect absorption by phagocytes. 
Loosanoff and Davis (1963) reviewed work done on the rearing of 
bivalve mollusks and summarized the effects of different foods on 
their growth. They developed a system of feeding oysters di- 
atoms and other algae. Also, they have made studies indicating 
that bivalve larvae can utilize dried algae. Coe (1948) indicates 
that carbohydrates are digested in the intestinal tract and the food 
is particulate in form. Guillard and Wangersky (1958) found that 
extra-cellular carbohydrates are produced by many flagellates. 
Rhamosides and ascorbic acids have been isolated from sea water 
by Wangersky (1952). Dexter Haven (personal communication) 
has succeeded in conditioning oysters by using suspensions of steri- 
lized cornstarch as a supplement in flowing sea water. 

Collier and co-workers (1953) made extensive studies on soluble 
and naturally occurring carbohydrates and how they affect oysters. 
They found that natural occurring carbohydrates will increase 
the pumping rate of oysters at levels as low as 2.25 mg per liter. 
Glutamate, glycogen, methionine, and inositol will cause the cirri 
of oysters to beat more rapidly. Collier (1959) also reported data 
which indicate that oysters can utilize dissolved carbohydrates for 
energy. Through a series of tests in which he compared the ca- 
loric intake of the oyster against caloric output of energy, he 



280 Quarterly Journal of the Florida Academy of Sciences 

found that the output of energy was greater than the caloric in- 
take. The output was measured by oxygen consumption; the 
intake was measured by the caloric value of the algae used for 
food. 

Experimental Procedure 

Two experiments, one with oysters and one without oysters, 
constituted the study. The first experiment was designed to com- 
pare the longevity of oysters on starvation diet against the length 
of life of those receiving only glucose. The second experiment 
was designed to compare the concentration of glucose in the 
tanks with the oysters, from the first experiment, as against the 
concentration of glucose in tanks under similar conditions but 
without oysters. 

Experiment Number 1 

Two epoxy-covered plywood tank systems that contained 310 
liters of water each when maintained at the experimental level 
were used. The water was circulated in the tanks, passed through 
filters, and aerated by means of baffles. The filters each contained 
one pound of glass wool and five pounds of activated animal char- 
coal, and the water was filtered four hours per day. The pur- 
pose of the filters was to remove some of the foreign particles, 
metabolites, bacteria and other substances that might be produced 
in this system. Artificial sea water (Lyman and Fleming formula) 
adjusted to a range of 17.5 to 20 o/oo was used. The mixture was 
formulated and adjusted throughout the experiment with deion- 
ized water. NaHC0 3 was used to buffer the first batch to a pH 
of 7.9; in all other batches, Tris buffer was used to maintain a 
pH of 7.9 to 8.1. The water was changed approximately every 
four weeks. 

Twenty-five scrubbed three-inch local oysters, Crassostrea vir- 
ginica (Gmelin), were added to each tank. The control tank (No. 
1), did not receive any glucose, and the experimental tank (No. 2) 
was fed as needed at a rate of 1.5 to 3.1 grams per day. An at- 
tempt was made to keep the concentration of the glucose in the 
tank between 5 and 15 mg per liter of water. 

The total carbohydrate content was checked periodically using 
the anthrone method as described by Lewis and Rakestraw (1955). 
The mortalities, temperature and salinities were recorded daily, 
and the pumping of the oyster as indicated by gaping was checked 



Gillespie, Ingle, Havens: Nutrition in Oysters 281 

three times daily. It is recognized that gaping shells do not always 
prove that oysters are pumping. However, the correlation be- 
tween the two events is generally fair and in a very gross way can 
be used as an index of pumping. It might also be argued that 
closed shells definitely preclude pumping; thus, pumping would 
be restricted to open valved individuals. 

Experiment Number 2 

The same type of tank system, filter, and carbohydrate test as 
explained in Experiment 1 were used, but no oysters were intro- 
duced. For identification the tanks were labeled No. 3 and No. 4. 
Tank 3 was not filtered and Tank 4 was filtered four hours daily. 
The water pH and salinity were also maintained at the same level 
as in the previous experiment, and an attempt was made to main- 
tain a maximum concentration of 15 mg per liter of water in each 
tank. The study was carried on for 24 days but no tests were made 
on the week ends. 

Results and Discussion 

A comparison of the concentration of carbohydrates for both 
experiments is given in Table 1. 

The steps of activities relating to glucose addition, filtration 
and related procedure are given here in order of their routine se- 
quence. 

1. Samples taken for the an throne test. 

2. Filtration. 

3. Samples taken for the an throne test. 

4. Glucose added. 

5. Samples taken occasionally for the anthrone test. 

The total glucose in the tanks was established by adding the re- 
sults of steps 3 and 4. The total glucose removed was determined 
by subtracting the results from step 3 from the total glucose found 
the previous day. In a similar manner the glucose removed by 
the filter was calculated by subtracting the results of step 3 from 
step 1. Also, the glucose removed by factors other than filter was 
determined as the difference between the total glucose removed 
and the glucose removed by the filter. All calculations are based 
on average daily values. Step 5 served only as a check. 



282 Quarterly Journal of the Florida Academy of Sciences 

TABLE 1 

Average daily glucose concentration in tanks* 



Amount of glucose 


Tank 1 


Tank 2 


Tank 3 


Tank 4 


Concentration (mg/1) 


.84 


13.00 


11.19 


11.06 


Grams added to maintain 










concentration 


— 


2.43 


.86 


2.11 


Total grams in tank 


.26 


4.03 


3.47 


3.43 


Total grams removed from tank 


.01 


2.55 


1.11 


2.35 


Grams removed by filter 


— 


.29 


— 


.70 


Grams removed by factors 










other than filter 


— 


2.26 


— 


1.64 


Per cent of total glucose 










removed from tank 


— 


63.22 


31.82 


67.10 


Per cent of glucose removed 










by filter 


— 


11.20 


— 


30.62 


Percent of glucose removed by 










factors other than filter 


— 


88.80 


— 


69.38 



*Total carbohydrate is recorded as glucose. 

Tank 1: 85 days on test; with oysters; no glucose added; filtered 4 
hours daily. 

Tank 2: 69 days on test; with oysters; glucose added; filtered 4 hours 
daily. 

Tank 3: 16 days on test; without oysters; glucose added; unfiltered. 

Tank 4: 16 days on test; without oysters; glucose added; filtered 4 
hours daily. 

The anthrone test was taken periodically and the number of 
days recorded in this table were selected from the days that the 
test was run. An occasional test showed a slight increase in glucose 
concentration after filtration. These tests were disregarded; how- 
ever, they would not have changed the results significantly. 

Tank 1, which did not receive any glucose, had an average 
daily concentration of 0.84 mg/liter. Tank 2 had a maximum daily 
average of 13.00 mg/liter, which was equivalent to 4.03 gm of 
glucose in the system. An average of 2.26 gm of glucose was 
removed from the tank daily, in the sense that the glucose 
could no longer be detected as carbohydrate. Of the amount re- 
moved 11.20 per cent was removed by the filter and 88.80 per cent 
by other factors. 



Gillespie, Ingle, Havens: Nutrition in Oysters 



283 





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400 



200 
DAYS 

Fig. 1. Longevity of starved and glucose-fed oysters. 



In the experiment without oysters, the non-filtered tank (No. 3) 
had a daily maximum concentration of 11.19 mg/liter which is 
equivalent to 3.47 gm in the tank. Of this, 31.82 per cent or 1.11 
gm was removed daily. In the filtered tank (No. 4), the average 
daily maximum level was 11.06 mg /liter or 3.4 gm in the tank. 
Of the glucose removed, 69.38 per cent was removed by factors 
other than the filter. In comparing Tank 3 with Tank 4, it be- 
came apparent that filtration accounts for over 30 per cent of the 
glucose removed. 

In comparing Tank 2 with Tank 4, 0.32 gm more glucose was 
needed daily to maintain the concentration in the tank with oys- 
ters. Furthermore, in the tanks with the oysters, 88.80 per cent 
of the glucose removed was removed by factors other than the 
filter; in the tank without oysters 69.38 per cent was similarly re- 



284 Quarterly Journal of the Florida Academy of Sciences 

moved. This would indicate that about 19.5 per cent of the glucose 
removed was used by the oysters. This loss of glucose could be 
accounted for either by direct assimilation or by an intermediate 
step of oysters feeding on micro-organisms which in turn had 
utilized the carbohydrate. 

A comparison of the longevity of the oysters in Tanks 1 and 2 
for one year is given in Fig. 1. At the end of 12 days, two oysters 
were sacrificed from each tank for a glycogen test and they were 
not recorded in this data. 

The number of deaths for tanks No. 1 and 2, respectively, were 
as follows: 50 days— 1, 0; 100 days— 2, 0; 150 days— 6, 1; 200 
days— 17, 5; 250 days— 19, 13; 300 days— 22, 17; 350 days— 22, 18. 
At the end of 390 days all oysters had died in Tank 1 and 19 in 
Tank 2. On the basis of the mortality at the end of one year, it 
was found that oysters being fed glucose lived on an average of 
68.2 days longer than the oysters not being fed. 




Fig. 2. Photograph of oysters at end of experiment. Left: Last sur- 
viving starved oyster after 390 days in artificial sea water. Right: One of 
four typical oysters surviving after 390 days in artificial sea water plus glucose. 



Gillespie, Ingle, Havens: Nutrition in Oysters 285 

During the 390th day of the experiment, the last oyster which 
had died from Tank 1 and the four oysters from Tank 2 were 
opened and examined. Figs. 2-3 compare the condition of the last 
oyster from Tank 1 with a typical individual from Tank 2. Extreme 
emaciation with almost complete degeneration of the mantle was 
noted in the starved oyster. Lack of "fatness" and a much smaller 
degree of emaciation were observed in the oysters from Tank 2. 
In the authors' opinion, the conditions noted occurred before death 
and were not due to post-mortem degeneration of tissue. 





Fig. 3. Interpretative outline drawing of oyster meats derived from pho- 
tograph in figure 2. Left: Meat of oyster starved 390 days. Right: Meat of 
oyster glucose-fed for 390 days. 

It is recognized that glucose by itself is not a complete diet and 
would not be expected to sustain the oyster completely. It is of 
interest to note that the oyster when not subject to predation, dis- 
ease or exorbitantly rigorous ecological conditions is very hardy 
and even when starved it is capable of living over a year. 

The tenacity for life shown by the starved animals emphasizes 
anew the importance of infections, large carnivores, and extremes 
of hydrographic conditions in the mortalities of oysters. A lack 
of importance is suggested for temporary periods of nutritional 
inadequacies. This lack of importance has been suggested in 
earlier work. Mackin (personal communication) advises that in 
connection with other studies, he kept oysters alive in filtered 
water, from which presumably all organism and detritus were 
removed, for periods of many months. In a separate discussion of 
mortality, Mackin (1961) postulates a scant possibility for natural 
starvation. 

The oyster tanks were housed in a concrete shed with screened 
doors and windows on all sides. They were always in the shade 



286 Quarterly Journal of the Florida Academy of Sciences 

and out of direct weather but otherwise subjected to ambient 
atmospheric variations. The daily temperatures listed as biweekly 
averages are given in Fig. 4. Since there was little or no variation 
in the temperature between tanks, the data are given as averages 
of the two tanks. Tanks 3 and 4 were housed in the same shed, 
and the water temperatures were essentially the same as given 
in this chart for the period covered. The temperature varied from 
a low of 1.8 °C on one occasion to a high of 28.0 °C on four days. 
For fifteen consecutive weeks, the daily average water tempera- 
ture was above 25 °C. Of this, four weeks averaged above 27 °C 
and nine weeks averaged above 26° C. Seventeen oysters died 
through the 19th to 25th weeks inclusive. During this seven week 
period, the average water temperature was 26 °C or above. It 
could not be said with certainty that the high temperature accele- 
rated the death rate, but it appears that this was the case. It is 
of interest to note that of the seventeen oysters that died during this 
warm period only four were from the tank being fed glucose. 



30-, 



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5 10 15 20 25 30 35 40 45 50 55 

APRIL MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC. JAN. FEB. MARCH APRIL 

WEEKS 



Fig. 4. Biweekly average daily ambient temperatures. 



Gillespie, Ingle, Havens: Nutrition in Oysters 287 

Observations were made on the pumping of the oyster and it 
appears that glucose under the conditions of this experiment will 
stimulate the oyster to pump. Oysters that were not fed were 
very rarely observed with shells agape. 

Summary and Conclusions 

Two experiments were conducted to investigate the use of 
glucose in the nutrition of the oyster, Comparisons were made 
on the concentration and removal of glucose from tanks with and 
without oysters; and a study was also made to compare the lon- 
gevity of oysters starved with the length of life of oysters being 
fed glucose. The concentration of glucose in the water was main- 
tained within a range of 5 to 15 mg per liter. 

Under the conditions of these experiments, it was found that 
more glucose was needed to maintain the concentration in the 
tank containing oysters than in tanks without oysters. Also, evi- 
dence was developed to show that about 19.5 per cent of the glu- 
cose removed from the tank was utilized by the oyster; however, 
these data do not necessarily indicate direct assimilation. The 
oysters being fed glucose lived an average of 68.2 days longer 
than those which were starved. It was observed that when main- 
tained in open tanks and in artificial sea water that oysters are 
capable, in some instances, of living longer than one year and 
three weeks without being fed. 

More investigations are needed to ascertain whether oysters 
can directly utilize glucose or other soluble carbohydrates. Studies 
are also needed to determine if soluble carbohydrates can be fed 
as a supplement for growth with other nutrients or in natural non- 
filtered bay water. We are encouraged to pursue these experiments 
by the findings of Stephens (1961, 1962, 1963, 1964), whose work 
showed the uptake of labelled glucose and amino acids by several 
marine invertebrates. 

Literature Cited 

Coe, Wesley R. 1948. Nutrition environmental conditions and growth of 
marine bivalve mollusks. Jour. Marine Research, vol. 7, no. 3, pp. 
586-601. 

Collier, Albert. 1959. Some observations on the respiration of American 
oyster Crassostrea virginica (Gmelin). Inst. Marine Sci., vol. 6, pp. 
92-108. 



288 Quarterly Journal of the Florida Academy of Sciences 

Collier, A., S. M. Ray, A. W. Macnitzky, and J. Bell. 1953. Effects of 
dissolved organic substances on oysters. U. S. Dept. of Interior, Fish- 
ery Bull., no. 84, pp. 167-185. 

Guillard, R. R. L., and P. J. Wagnersky. 1958. The production of extra- 
cellular carbohydrates by some marine flagellates. Limnology and 
Oceanography, vol. 3, no. 4, pp. 449-454. 

Lewis, G. J., Jr., and N. W. Rakestraw. 1955. Carbohydrate in sea water. 
Jour. Marine Research, vol. 14, no. 3, pp. 253-258. 

Loosanoff, V. L., and H. C. Davis. 1963. Rearing of bivalve mollusks. 
Advances in Marine Biology. Academic Press, New York, pp. 1-136. 

Mackin, J. G. 1961. Mortalities of oysters. Proc. Nat. Shellfish Assoc, vol. 
50, pp. 21-40. 

Stephens, Grover C, and R. A. Schinske. 1961. Uptake of amino acids by 
marine invertebrates. Limnology and Oceanography, vol. 6, no. 2, pp. 
175-181. 

Stephens, Grover C. 1962. Uptake of organic material by aquatic inverte- 
brates. I. Uptake of glucose by the solitary coral, Fungia scutaria. 
Biol. Bull, vol. 123, no. 3, p. 648-659. 

. 1963. Uptake of organic material by aquatic invertebrates. II. 

Accumulation of amino acids by the bamboo worm, Clymenella torqua- 
ta. Comp. Biochem. Physiol., vol. 10, pp. 191-202. 

. 1964. Uptake of organic material by aquatic invertebrates. III. 

Uptake of glycine by brackish-water annelids. Biol. Bull., vol. 126, 
no. 1, pp. 150-162. 

Wagnersky, P. J. 1952. Isolation of ascorbic acid and rhamnosides from 
sea water. Science, vol. 115, p. 685. 

Yonge, C. M. 1928. The absorption of glucose by Ostreaedulis. Jour. Ma- 
rine Biol. Assoc, vol. 15, pp. 643-653. 

Florida Board of Conservation Marine Laboratory, St. Peters- 
burg, Florida. Contribution No. 82. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



NOTES ON POSTLARVAE OF PANULIRUS ARGUS 
Ross Witham, Robert M. Ingle, and Harold W. Sims, Jr. 

In June of 1963 a random push net sample taken in the Indian 
River adjacent to the dock of the House of Refuge Museum near 
Stuart, Florida, produced several postlarval specimens of the spiny 
lobster Panulirus argus (Latr.). The existence of adult populations 
of P. argus in the offshore waters of the Martin County area has 
been reported by Moore (1962) and Robinson and Dimitriou (1963). 
To our knowledge this is the first report of postlarval spiny lobster 
in this area. 

Literature on the ecology of the early postlarval stages of 
Panulirus argus is nonexistent and the study of the growth rate is 
limited to the work of Lewis et al. (1952). The lack of data on 
this phase of lobster research and the established availability of 
postlarvae in the Stuart area led us to begin this study. Postlarvae 
were collected at random in the waters of the Indian River, the 
Intracoastal Waterway south of the St. Lucie Inlet, and in the 
waters of Hobe Sound. This report provides preliminary data ob- 
tained during the first year of work. 

Methods and Materials 

Samples were taken using a variation of the push net commonly 
used in the collection of invertebrates in grass flats. The frame 
of the net was made in the form of a 3 ft by 3 ft square using one 
inch Polyvinyl chloride tube. The bag of the net was built as a 
shallow pocket formed of fiberglass screen. 

The method of collecting varied with the substrate. In man- 
groves (Rhizophora sp.) the net was placed under a mass of algae 
or other organisms growing on the submerged roots; with the net 
in place, the attached organisms were pulled from the.rOots and 
allowed to fall into the bag. On grass flats the net was inserted 
under a mass of unattached plants and lifted; the algae thus col- 
lected were examined for postlarvae. On seawalls, piling, and rock, 
where the sessile plants could not be removed by the former meth- 
ods, a % meter plankton net was fastened to an 8 foot pole, with 
a metal scraper attached above the net; this scraped the growths 
from the solid submerged object, with the sample falling in the 
plankton net. In rocky areas two small screen nets were used; 



290 Quarterly Journal of the Florida Academy of Sciences 

the sample was gripped between the nets, pulled loose, and brought 
to the surface for examination. 

Some of the postlarvae were taken to the laboratory for feeding 
and growth observations. Others were preserved in 10 per cent 
formaldehyde. 




Map 1. Collecting stations near Stuart, Florida. 



Witham, Incle, Sims: Postlarvae of Panulirus 291 

General Description of the Area 

The inland waters of this area can best be described as two 
shallow lagoons formed by offshore barrier islands (Map 1). They 
are connected by the Intracoastal Waterway, although historically 
a shallow, tortuous meander joined them. The water depth, except 
in the channels, is relatively shallow, most of the areas being less 
than a fathom at low tide. Dense beds of Syringodium and Di- 
planthera are common throughout the area except in the Intra- 
coastal Waterway and in Hobe Sound, where they are very sparse 
(Phillips and Ingle, 1960). 

Marine organisms, principally plants, found throughout the 
area in which the postlarvae were found, varied in density through- 
out the year. Included were such species as Bugula sp., Giffordia 
mitchellae (Harv.), Gracilaris cylindrica (B0rgs.), Acanthophora 
spicifera (Vahl.), Hypnea cervicornis (J. Ag.), Hypnea cornuta 
(Lamx.), Erythrotrichia carnea (Dillw.), Ceramium brevizonatum 
H. E. Peterson, and Acroochaetium sargassii (B0rgs.). These were 
listed by Phillips (personal communication) and were found at- 
tached or unattached. Stations 1 and 3-10 are located in typical 
shallow grass beds with mangrove-lined shores. Station 2, located 
at Seminole Shores Marina (S. S. on chart), has seawalls, pilings, 
rocks, and some small grass flats. 

Station A is located on the mangrove shore line bordering the 
north side of the St. Lucie Inlet. Stations S1-S5 are situated in 
the Intracoastal Waterway and Hobe Sound area. The 16 stations 
shown on Map 1 were sampled at random during the 13 month 
period of this study. Station data obtained are provided in Table 1. 

Results ~ 

Our data indicate that the transparent puerulus stage begins 
to appear in small numbers during the months of January and 
May and becomes increasingly common throughout the summer 
and fall, the peak number being reached during October and No- 
vember. These findings do not coincide with those of Lewis et al. 
(1952), who worked in the Miami area and found January to be the 
most productive for this stage. No explanation is offered at this 
time for this discrepancy. A comparison of January water tempera- 
tures for the two bays would possibly provide a clue. 



292 Quarterly Journal of the Florida Academy of Sclences 



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Witham, Ingle, Sims: Postlarvae of Panulirus 295 

Limited growth studies were made on two slightly advanced 
postlarvae held in separate tanks at our Stuart Laboratory. All 
measurements listed were made from the horns to the rear edge of 
the carapace, along the mid-dorsal line. 

Spiny lobster A was kept in a tank with continuously changing 
sea water and was fed a diet of dry cat food, "Purina Cat Chow" 
and "Kitty Krumpets." Measurements were made from the cast 
carapace exoskeleton after each molt. The first cast, which meas- 
ured 15 mm, was produced February 8; the second on March 24, 
1964, was 16 mm. On April 16 the third molt took place, and the 
carapace measured 17.5 mm. Molt number four occurred on May 
19, and a carapace of 21.5 mm resulted. The last molt was ob- 
served on July 26. The cast carapace measured 25 mm. Lobster 
A is still living at the time of this report. 

The carapace of spiny lobster B measured 32.5 mm at the time 
of the first molt in February. A second molt took place on April 
27, the cast carapace measuring 38 mm. On June 9 the third molt 
occurred, the cast measuring 45 mm. Molt number 4 was on July 
17. The carapace then measured 52 mm. This lobster was found 
dead on July 28, at which time the carapace length was 61 mm. 
Throughout the period of confinement this lobster was offered a 
diet of sea slugs, sea urchins, and fish but was observed to feed 
only on the latter. 

Other studies (Crawford and DeSmidt, 1922; Smith, 1948, 1950; 
Smith and Marshall, 1945; Dawson and Idyll, 1951; Sutcliffe, 1957) 
show a longer period between, and a smaller increase after, ob- 
served molts. These previous studies were, however, made on 
larger, more mature animals, so our more accelerated growth is 
not unexpected. Lewis's findings, on sizes similar to ours, gave 
results more in agreement with those we obtained. 

In addition to our postlaval studies several plankton samples 
were taken. Two phyllosoma larvae were taken in the St. Lucie 
Inlet on November 11, 1963. Samples taken in the Indian River 
near the House of Refuge on June 16 and 25, 1964, each had one 
first stage larva. These phyllosomes compare with the first stage 
of Panulirus argus as described by Lewis (1951). 

Recent work on estuarine biology has indicated a remarkable 
number of marine animals that appear to be obligated to spend 
their juvenile stages in the protected sanctuaries of river mouths 
and bays. Evidence presented here supports the belief that the 



296 Quarterly Journal of the Florida Academy of Sciences 

spiny lobster, sublittoral as an adult, may be added to the growing 
list of animals that are to some extent dependent upon sequestered 
shallows for juvenile development. 

Summary 

Postlarvae of the spiny lobster, Panulirus argus, are found in 
the Indian River north of the St. Lucie Inlet and the Intracoastal 
Waterway and Hobe Sound south of the St. Lucie Inlet. Phyllo- 
soma larvae have been taken in the St. Lucie Inlet and Indian 
River. Limited studies indicate that P. argus has a rapid rate of 
growth in early stages. Data obtained suggest that Florida's spiny 
lobster may be yet another marine animal utilizing estuaries and 
bays as sanctuaries and nurseries for juvenile stages. 

Literature Cited 

Crawford, D. R., and W. J. J. DeSmidt. 1922. The spiny lobster, Pan- 
ulirus argus, of southern Florida: its natural history and utilization. 
Bull. U. S. Bur. Fish., vol. 38, pp. 281-310. 

Dawson, C. E., Jr., and C. P. Idyll. 1951. Investigations on the Florida 
spiny lobster, Panulirus argus (Latreille). Florida State Board of Conser., 
Tech. Ser., no. 2, pp. 1-39. 

Lewis, J. B. 1951. The phyllosoma larvae of the spiny lobster Panulirus 
argus. Bull. Mar. Sci. Gulf and Carib., vol. 27, no. 2, pp. 89-103. 

Lewis, J. B., H. B. Moore, and W. Bahis. 1952. Post-larval stages of the 
spiny lobster Panulirus argus. Bull. Mar. Sci. Gulf and Carib., vol. 2, 
no. 1, pp. 324-337. 

Moore, D. R. 1962. Notes on the distribution of the spiny lobster Pan- 
ulirus in Florida and the Gulf of Mexico. Crustaceana, vol. 3, no. 4, 
pp. 318-319. 



PhillipsJ R. C, and R. M. Incle. 1960. Report on the marine plants, bot- 
tom types and hydrography of the St. Lucie estuary and adjacent Indian 
River, Florida. Florida State Board of Conser., Spec. Sci. Rept., no. 4, 
pp. 1-78. 

Rorinson, R. K., and D. E. Dimitriou. 1963. The status of the Florida 
spring lobster fishery, 1962-63. Florida State Board of Conser., Tech. 
Ser., no. 42, pp. 1-30. 

Smith, F. G. Walton. 1948. The spiny lobster industry of the Caribbean 
and Florida. Carib. Res. Coun. Carib. Comm. Port-of-Spain, Trinidad., 
Fish Ser., no. 3., pp. 1-49. 



Witham, Ingle. Sims: Postlarvae of Panulirus 297 

. 1950. Caribbean spiny lobster investigations. Proc. Gulf and 

Carib. Fish. Inst., Third Annual Session 1950, pp. 128-134. 

Smith, F. G. Walton, and N. Marshall. 1945. Preliminary report on the 
Florida crawfish investigation. Univ. Miami Mar. Lab., pp. 1-51. 

Sutcliffe, W. H., Jr. 1957. Observations on the growth rate of the imma- 
ture Bermuda spiny lobster, Panulirus argus. Ecology, vol. 38, no. 3., 
pp. 526-529. 

Florida Board of Conservation Field Station, Stuart, Florida. 
St. Petersburg Marine Laboratory Contribution No. 83. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



TWO DRAGONFLIES NEW TO FLORIDA 
Dennis R. Paulson 

Pending publication of a survey of the Odonata of southern 
Florida, I take this opportunity to put on record two species previ- 
ously unrecorded in the state. The first of these, Brachymesia 
herbida (Gundlach), has been taken at a brackish ditch on Big Pine 
Key, Monroe County, where adults were collected on 14 October 
1960 and 9 April 1961. Needham (1945) casually mentioned col- 
lecting this species at Clewiston, Hendry County, the only record 
of its occurrence in Florida. M. J. Westfall, Jr., however, has in- 
formed me (in litt., 27 March 1961) that Needham's record is based 
upon immature specimens of Brachymesia gravida (Calvert). 

The second species, N eoerythromma cultellatum (Hagen), has 
been taken at Nine Mile Pond in Everglades National Park, Dade 
County, where adults were collected 9 July 1962 and 13 June 1963, 
and an emerging individual was found on 15 March 1964. The 
species is apparently established as a resident. 

Specimens of both species are presently in my possession; rep- 
resentatives are to be deposited in the University of Florida col- 
lections. 

Both species are widespread in the Greater Antilles. B. herbida 
also ranges from southern Texas to Bolivia and Brazil. N. cultella- 
tum has been recorded from Guatemala to Panama, and may be 
more widespread than that on the mainland, as I have examined 
two males in the University of Florida collections from La Gloria 
Cardel, Veracruz, Mexico, collected in 1938 by J. Camelo G. The 
present record is the first for N. cultellatum in the United States. 
The finding of these two species in southern Florida further 
strengthens the idea of this area as a transitional one between tem- 
perate and tropical faunas. 

Literature Cited 

Needham, J. G. 1945. Notes on some dragonflies of southwest peninsular 
Florida. Bull. Brooklyn Ent. Soc, vol. 40, pp. 104-110. 

Department of Zoology, University of Miami, Coral Gables, 
Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965): 298 



NEW ATLANTIC COAST RANGES FOR FISHES 

William D. Anderson, Jr., and Elmer J. Gutherz 

Specimens of Corniger spinosus, Holocentrus ascensionis, Myri- 
pristis jacobus, and Ostichthys trachypomus (Holocentridae); Ani- 
sotremus virginicus (Pomadasyidae); Chaetodon sedentarius and 
Holacanthus tricolor (Chaetodontidae); and Bodianus pulchellus 
and Clepticus parrai (Labridae) have been collected by the U. S. 
Bureau of Commercial Fisheries exploratory fishing vessels Combat, 
Silver Bay, and Delaware at localities along the Atlantic coast of 
the United States north of their previously reported ranges. These 
specimens are in the collections of the U. S. Bureau of Commercial 
Fisheries Biological Laboratory, Brunswick, Georgia, unless other- 
wise indicated. We are grateful to Loren P. Woods of the Chicago 
Natural History Museum and Bruce B. Collette of the Bureau of 
Commercial Fisheries Ichthyological Laboratory, for their com- 
ments on the manuscript, and to Robert H. Gibbs, Jr., Division of 
Fishes, U. S. National Museum, for the loan of specimens. 

1. Corniger spinosus Agassiz 

Four specimens (103-122 mm S.L.) from Silver Bay stations 
1393, 2989, 5507, and 5579. Range previously reported as "costas 
del Brasii" and Cuba north of Havana by Howell Rivero (1941, pp. 
3-7). The collection of the specimen off Cape Remain, S. C. (Silver 
Bay station 1393), represents a range extension of about 650 nauti- 
cal miles northward from Cuba, along the Atlantic coast of the 
United States. 

2. Holocentrus ascensionis (Osbeck). Squirrelfish 

Twelve specimens (180-276 mm S.L.) from Silver Bay stations 
1534, 2999, 3000, 3330, 3649, 4672, 5417, 5418, 5419, and 5420; and 
four specimens (22-40 mm S.L., deposited in the U. S. National 
Museum) collected by Robert H. Gibbs, Jr. with dip net and night 
light, while aboard the Delaware 38°56'N. Lat., 66°27'W. Long. 
(USNM 194241), 39°07'N. Lat., 65°58'W. Long. (USNM 194239), 
37°02 , N. Lat., 71°20'W. Long. (USNM 194240), and 39°44.5'N. 
Lat., 70°56'W. Long. (USNM 194244). Range previously reported 
as "Both side of the Atlantic; in the western Atlantic from Ber- 



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302 Quarterly Journal of the Florida Academy of Sciences 

muda and Florida to Rio de Janeiro and the northern and western 
Gulf of Mexico" by Briggs (1958, p. 271). McKenney (1959, p. 208) 
cited several obscure references in which Holocentrus had been 
reported from as far north as Newport, Rhode Island, and Woods 
Hole, Massachusetts; these records might have been of pelagic 
juveniles and might refer to H. ascensionis. Gibbs and Collette 
(1959, pp. 145-146) found four specimens of H. ascensionis in the 
stomach contents of a Coryphaena hippurus "from the Gulf Stream." 
The stomach contents mentioned by Gibbs and Collette were col- 
lected from the Delaware at 38°58'N. Lat, 66°26'W. Long, on 
9 June 1957 (Bruce B. Collette, in litt., 19 August 1964). The col- 
lection of the specimen (USNM 194244) from SE of Montauk Pt, 
Long Island, N. Y., represents a range extension of about 800 nauti- 
cal miles northward from Florida along the Atlantic coast of the 
United States. 

3. Myripristis jacobus Cuvier. Blackbar soldierfish 

Three specimens (101-142 mm S.L.) from Combat station 353 
and Silver Bay station 3009. Range previously reported as "Both 
sides of the Atlantic; in the western Atlantic from the northeastern 
Gulf of Mexico to Tortugas, Florida, and south to Rio de Janeiro" 
by Briggs (1958, p. 270). The collection of the specimen off Jack- 
sonville Beach, Fla. (Combat station 353), represents a range ex- 
tension of about 500 miles by sea from Tortugas. 

4. Ostichthys trachypomus (Giinther). Bigeye soldierfish 

One specimen (29 mm S.L.) from Silver Bay station 4117 and 
three specimens (22-25 mm S.L., deposited in the U. S. National 
Museum) collected by Robert H. Gibbs, Jr. with dipnet and night 
light, while aboard the Delaware 38°05'N. Lat, 65°58'W. Long. 
(USNM 194245) and 39°44.5'N. Lat., 70°56'W. Long (USNM 
194246). Range previously reported as West Indies and Cuba by 
Jordan, Evermann, and Clark (1930, p. 234). Bailey et al. (1960, 
p. 24) first listed this species as occurring in waters of the United 
States presumably based upon a previously unpublished record 
off North Carolina furnished by Loren P. Woods. The collection 
of the specimens (USNM 194246) from SE of Montauk Pt., Long 
Island, N. Y., represents a range extension of about 900 nautical 
miles northward from the northernmost part of the West Indies 
(Bahamas), along the Atlantic coast of the United States. 



Anderson and Gutherz: New Fish Ranges 303 

5. Anisotremns virginicus (Linnaeus). Porkfish 

Three specimens (82-260 mm S.L.) from Silver Bay stations 
5516 and 5682, Range previously reported as "Bermuda and south- 
ern Florida to Santa Catarina, Brazil, and the eastern and southern 
Gulf of Mexico" by Briggs (1958, p. 279). The collection of the 
specimens off Anastasia Is., Fla. (Silver Bay station 5682), repre- 
sents a range extension of about 200 nautical miles northward 
from southern Florida along the Atlantic coast of the United 
States. 

6. Chaetodon sedentarius Poey. Reef butterflyfish 

Thirty-seven specimens (73-120 mm S.L.) from Silver Bay sta- 
tions 1207, 1233, 1268, 1297, 1506, 1534, 1734, 1738, 2543, 3010, 
4652, 4665, 4673, 4921, 4938, 5390, 5391, 5393, 5417, 5420, and 
5437. Range previously reported as Bermuda and West Indies by 
Beebe and Tee-Van (1933, p. 176); as "Eastern and southwestern 
Gulf of Mexico to Hispaniola" by Briggs (1958, p. 282); from the 
northern Gulf of Mexico by Collins and Smith (1959, p. 252); and 
from off the south Atlantic coast of the United States, but with no 
specific locality given, by Hubbs (1963, p. 140). The collection of 
the specimen off Cape Lookout, N. C. (Silver Bay station 1268), 
represents the northernmost known occurrence of this species along 
the Atlantic coast of the United States. 

7. Holacanthus tricolor (Bloch). Rock beauty 

Four specimens (115-173 mm S.L.) from Silver Bay stations 
5418, 5419, and 5420. Range previously reported as "Bermuda and 
the Florida Keys to Rio de Janeiro" by Briggs (1958, p. 283). The 
collections of these specimens off Cumberland Is., Ga., represent 
a range extension of about 300 nautical miles northward along the 
Atlantic coast of the United States. 

8. Bodianus pidchell us. (Poey). Spotfin hogfish 

Five specimens (176-231 mm S.L.) from Silver Bay stations 
4665, 4673, 5417, and 5418. The specimen from Silver Bay 4665 
is deposited in the Chicago Natural History Museum (CNHM 
66683). Range previously reported as "southeastern Florida and 
the Bahamas to northern South America" by Feddern (1963, p. 



304 Quarterly Journal of the Florida Academy of Sciences 

227). The collections of the specimens off Cape Romain, S. C. 
(Silver Bay stations 4665 and 4673), represent a range extension of 
about 450 nautical miles northward of that given by Feddern. The 
largest specimen examined by Feddern was "156.2" mm (UMML 
9508). All specimens collected by the Silver Bay were larger but 
show the total gillraker count given by Feddern to distinguish 
Bodianus pulchellas (15 to 16) from the closely related Bodianus 
rufus (17 to 19). The preserved specimens show distinctive mark- 
ings similar to those described by Feddern for B. pulchellus (a 
striking black area on the dorsal part of the pectoral distally and 
a black spot at the bases of the first two to four dorsal spines). 

9. Clepticus parrai (Bloch and Schneider). Creole wrasse 

One specimen (37 mm S.L.) from Silver Bay station 4183. Range 
previously reported as Bermuda and the West Indies by Beebe and 
Tee- Van (1933, p. 194). Henry A. Feddern (in litt., 16 March 1964) 
stated that there are "quite a few specimens of Clepticus parrai" 
in the collections of the University of Miami Marine Laboratory; 
all from the Florida Keys except one found dead on the beach at 
Miami Beach, Dade County, Florida, on 10 September 1960 
(UMML 7140). The collection of this specimen off Cape Fear, 
N. C, represents a range extension of about 550 nautical miles 
northward along the Atlantic coast of the United States. 

Discussion 

Captures of large Holocentrus ascensionis and Chaetodon se- 
dentarius from many localities off the south Atlantic coast of the 
United States in all seasons show they are resident species as far 
north as northern South Carolina (H. ascensionis) and central 
North Carolina (C. sedentarius), whereas the small specimens of 
H. ascensionis dipnetted farther north probably drifted into their 
areas of capture. 

Collections of Bodianus pulchellus in January and December 
1963 (nearly a year apart) and of Holacanthus tricolor in December 
1963 during a time of the year when we would not expect warmer 
water strays, and each at more than one locality, indicate that these 
species normally range this far north. 

Ostichthys trachypomus and Clepticus parrai may not normally 
occur as far north as these records suggest. The specimens of each 



Anderson and Gutherz: New Fish Ranges 305 

of these species could have developed from pelagic larval forms 
which drifted into their areas of capture. Their small size and 
capture in the warmer months (O. trachypomus in June, August, 
and September and C. parrai in July) lend credence to this hypoth- 
esis. 

Increased collecting in progress off the Atlantic coast of the 
United States will likely reveal that many species now considered 
sub-tropical or tropical have far more extensive northerly ranges — 
as strays or random wanderers, as migrants, and as indigenous 
members of the fauna. 

Literature Cited 

Bailey, Reeve M., et al. 1960. A list of common and scientific names of 
fishes from the United States and Canada. Am. Fish. Soc, Spec. Publ., 
no. 2, pp. 1-102. 

Beebe, William, and John Tee- Van. 1933. Field book of the shore fishes 
of Bermuda. G. P. Putnam's Sons, 337 pp., illus. 

Briggs, John C. 1958. A list of Florida fishes and their distribution. Bull. 
Florida State Mus., vol. 2, no. 8, pp. 223-318. 

Collins, Richard A., and Robert E. Smith. 1959. Occurrence of butterfly 
fish in Mississippi Sound. Copeia, no. 3, p. 252. 

Feddern, Henry A. 1963. Color pattern changes during growth of Bodi- 
anus pulchellus and B. rufus (Pisces: Labridae). Bull. Mar. Sci. Gulf 
and Caribbean, vol. 13, no. 2, pp. 224-241. 

Gibbs, Robert H., Jr., and Bruce B. Collette. 1959. On the identifica- 
tion, distribution, and biology of the dolphins, Coryphaena hippurus 
and C. equiselis. Bull. Mar. Sci. Gulf and Caribbean, vol. 9, no. 2, pp. 
117-152. 

Howell Rivero, Luis. 1941. "Corniger spinosus" Agassiz: nueva especie 
para Cuba y algunas consideraciones acerca de la misma. Torreia, no. 
6, pp. 3-7. 

Hubbs, Carl L. 1963. Chaetodon aya and related deep-living butterflyfish- 
es; their variation, distribution and synonymy. Bull. Mar. Sci. Gulf and 
Caribbean, vol. 13, no. 1, pp. 133-192. 

Jordan, David Starr, Barton Warren Evermann, and Howard Walton 
Clark. 1930. Check list of the fishes and fishlike vertebrates of 
North and Middle America north of the northern boundary of Vene- 
zuela and Colombia. Rept. U. S. Comm. Fish., 1928, App. X., pp. 
1-670. 



806 Quarterly Journal of the Florida Academy of Sciences 

McKenney, Thomas W. 1959. A contribution to the life history of the 
squirrel fish Holocentrus vexillarins Poey. Bull. Mar. Sci. Gulf and 
Caribbean, vol. 9, no. 2, pp. 174-221. 

U. S. Bureau of Commercial Fisheries Biological Laboratory, 
Brunswick, Georgia. Contribution number 76. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



HYPERTENSIVE EFFECT OF LATRODECTUS VENOMS 
John D. McCrone and Roger J. Porter 

The venoms of several subspecies of the black widow spider, 
Latrodectus madam, have a hypertensive effect on the mammalian 
systemic arterial pressure. Troise (1928), Sampayo (1944), Cicardo 
(1954) and Calvo et al. (1957) demonstrated this with the venom 
of L. mactans mactans, Shapiro et al. (1939) with the venom of L. 
mactans indistinctus, and Bettini and Toschi-Frontali (1960) with 
the venom of L. mactans tredecimguttatus. 

This paper presents the results of a study that was undertaken 
to determine whether the venoms of three other species of Latro- 
dectus, L. variolus, L. bishopi, and L. geometricus, have a similar 
hypertensive effect. This study is part of a more extensive investi- 
gation of the comparative toxicology and biochemistry of the ven- 
oms of the North American Latrodectus. 

Materials and Methods 

Lyophilized venom gland extract reconstituted with physiolog- 
ical saline was used in all experiments. The method of prepara- 
tion of this extract is given in a previous paper (McCrone, 1964). 
The glands were taken from mature female L. mactans mactans, 
L. variolus, L. bishopi, and L. geometricus collected in the state of 
Florida. 

The 2.1-3.8 kg rabbits used in this study were anesthetized with 
intravenous Nembutal (Abbot). The dose for each rabbit was de- 
termined individually by observing the eyelid reflex, the average 
being about 5-6 cc of a 12.5 mg/cc solution. The rabbits were 
then heparinized with intravenous Panheparin (Abbot). The femo- 
ral artery was exposed and cannulated with a small polyethylene 
tube which was connected to a recording mercury manometer by 
means of a hypodermic extension tube. Locke's solution contain- 
ing a small amount of the anticoagulant dye chlorazol fast pink was 
used in the connecting tube. After normal mean arterial pressure 
was established and maintained, various dosages of venom were 
injected into the ear veins of the rabbits and the changes in the 
mean arterial pressure were recorded on smoked kymograph paper. 



308 Quarterly Journal of the Florida Academy of Sciences 



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McCrone and Porter: Black Widow Venom 309 

Results 

A total of 12 experiments were performed, 3 with each venom. 
An increase in the mean systemic arterial pressure was observed 
in all cases; the increases ranged from 4-57 mm Hg beginning after 
a latent period of 27-140 seconds, and with one exception reaching 
a maximum after 1-16 minutes. A summary of the results of these 
experiments is given in Table 1. 

An unusual effect was recorded in rabbit number 6 which re- 
ceived 0.6 mg of L. variolus venom. In all other animals the max- 
imum pressure was reached in less than 16 minutes, but in this 
animal there was an initial small increase of 8 mm Hg after a 
latent period of 80 seconds, followed by a return to normal after 
2 minutes, with subsequent gradual increase in pressure until a 
maximum increase of 15 mm Hg was recorded after 41:50 minutes. 

Discussion 

Our studies demonstrate that all of the known North American 
species of the genus Latrodectus have venoms which have a hyper- 
tensive effect on the mammalian systemic arterial pressure. Mc- 
Crone (1964) has previously shown that these venoms are lethal to 
mammals. Troise (1928) and Sampayo (1944) on the basis of their 
experiments concluded that the hypertensive factor in the venom 
is separate from the lethal neurotoxic factor. Bettini and Toschi- 
Frontali (1960), however, found the same protein fraction is re- 
sponsible for both effects based on fractionation studies using 
paper electrophoresis. 

Acknowledgments 

We wish to thank Dr. Frederick Nichols for his helpful advice. 
This study was made possible by a grant from the Florida Heart 
Association and Public Health Service Grant GM 11206-01. 

Literature Cited 

Bettini, S., and N. Toschi-Frontali. 1960. Biochemical and toxicological 
aspects of Latrodectus tredecimguttatus venom. Verh. XI Internat. 
Kongr. Ent. Wien, vol. 3, pp. 115-121. 

Calvo, R., J. Chionetti, J. Fasciolo, A. Bania, M. Puebla, E. Zangheri, 
and F. Fernandez. 1957. Mecanismo de la accion presora del ven- 
eno de arana L. mactans. Rev. Soc. Argentina Biol., vol. 33, no. 678, 
pp. 309-319. 



310 Quarterly Journal of the Florida Academy of Sciences 

Cicardo, V. 1954. Mecanismo de la hipertension arterial producida par la 
ponzona de Latrodectus mactans. Rev. Soc. Argent. Biol., vol. 30, pp. 
19-24. 

McCrone, J. 1964. Comparative lethality of several Latrodectus venoms. 
Toxicon, vol. 2. (in press) 

Sampayo, R. 1944. Pharmacological action of the venom of Latrodectus 
mactans and other Latrodectus spiders. Jour. Pharmacol, and Exp. 
Therap., vol. 80, no. 4, pp. 309-322. 

Shapiro, H., N. Sapeika, and M. Finlayson. 1939. Pharmacological actions 
of the venom of Latrodectus indistinctus. S. African Jour. Med. Sci., 
vol. 4, pp. 10-17. 

Troise, E. 1928. Action pharmacodynamique du venin de Latrodectus mac- 
tans. Compt. Rend. Soc. Biol., vol. 99, no. 31, pp. 1431-1433. 

Florida Presbyterian College, St. Petersburg, Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



A NEW GLASS LIZARD FROM VERACRUZ, MEXICO 
J. Alan Holm an 

Our knowledge of Ophisaurus Daudin in Mexico is obscure. 
In fact, Smith and Taylor (1950, p. 194, footnote 97) emphatically 
excluded the genus from their Mexican checklist. McConkey (1954 
and 1955) established the presence of Ophisaurus in Mexico, but 
unfortunately his specimens were imperfect. A lizard with a 
crushed head from near Valles, San Luis Potosi, was named Ophi- 
saurus incomptus McConkey, and the remains of another specimen 
that had been hacked to death with a machete was tentatively 
assigned to the new species. The latter specimen was from Laguna 
de Los Cocos, Veracruz. A third specimen (USNM 6078) with the 
label "Jalapa, Veracruz" has been mentioned by Yarrow (1884) 
and Cope (1900), but McConkey (1954) has found it to be typical 
O. attenuatus attenuatus (a United States subspecies) and states 
"It is probably true that USNM 6078 has incorrect data". Smith 
and Taylor are more positive in their rejection of this record. 

During August of 1964 a collecting trip to Veracruz by the 
author and his wife yielded an additional Mexican specimen of 
Ophisaurus from coastal dune-scrub within the suburbs of the city 
of Veracruz. Fortunately, the specimen is almost perfect, although 
it does have a partially broken tail. This form exhibits character 
differences that are consistent with those previously found reliable 
in the definition of Ophisaurus species. I wish to call this new 
Veracruz specimen 

Ophisaurus ceroni sp. nov. 

Diagnosis. An Ophisaurus differing from other New World 
members of the genus by the following combination of charac- 
ters: frontonasal divided; scales along lateral fold 101; four indis- 
tinct, white, vertical bars on each side of neck; white spots on dor- 
sum absent; distinct, dark mid-dorsal stripe present; no dark pig- 
mentation below lateral fold. 

Holotype. Museum of Natural History Illinois State Univer- 
sity Number 272 (figs, la and b); taken by J. Alan and Donna Rae 
Holman from Veracruz, Veracruz, Mexico, August 11, 1964; habi- 
tat, coastal dune-scrub. 



312 Quarterly Journal of the Florida Academy of Sciences 























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Fig. 1. Pigmentation of M.N.H.I.S.U. 272, Ophisaurus ceroni sp. nov. 
A, dorsal view of head and anterior part of body. B, lateral view of anterior 
part of body. Drawings about twice natural size. 

Etymology. Named in honor of Senor Carlos Ceron of Cuautla- 
pan, Veracruz, in recognition of his dilligent and acute aid to her- 
petologists of the last three decades. 

Description of Holotype. Snout- vent length 143 mm.; tail par- 
tially broken; head width 8.8 mm., eye diameter 2.9 mm.; dorsal 
scales in 14 longitudinal series; scales around parietal 7-7, upper 
labials 11-11, preoculars 3-3, postnasals 2-2; scales around tail 18; 
scales along lateral fold 101. 

Frontonasal divided; labials separated from orbit by lorilabials 
and suboculars; prefrontals in broad contact; upper postnasal in 
contact with supracanthal row as well as with anteriormost canthal; 
anterior frontonasal separates postinternasals; five supraoculars; 
canthals extending to just anterior of middle of eye; frontal broad 
and rectangular posteriorly, somewhat pointed anteriorly, anterior 
end meeting fused prefrontals; interparietal broad anteriorly, taper- 
ing to a point posteriorly; occipital about as broad as interparietal 
at its greatest width; frontoparietal in contact with third and fourth 
supraoculars; first and second upper labials in contact with nasal. 

Body broader than high; dorsal scales keeled; ventrals smooth 
and flat. Ear opening oval, larger than round nostril. 

Dorsal ground color grayish-brown; interrupted by three dis- 
tinct dark stripes, two lateral and one mid-dorsal. Each lateral 
stripe confined to a little less than one scale row; mid-dorsal stripe 
occupying adjacent halves of two scale rows. Discrete white spots 



Holm an: New Glass Lizard 313 

lacking on both back and sides; no dark pigment below lateral fold. 
Head light grayish-brown; both top and sides of head speckled with 
dark spots; lower jaw with a few dark spots. Four indistinct white, 
vertical bars present on each side of neck. 

Discussion. An exhaustive study of New World Ophisaurus 
by McConkey (1954) has shown that the species of this genus are 
very similar in external morphology, yet they remain distinct in 
nature and may be distinguished from one another on the basis of 
combinations of external characters. Skeletal differences tend to 
confirm the validity of the recognized species of Ophisaurus 
(Auffenberg, 1955, Etheridge, 1960 and 1961, Holman, 1958, and 
Weigel, 1962). 

Ophisaurus ceroni has a combination of strong characters that 
separates it from other New World species. Some of the most im- 
portant of these characters follow: In O. ceroni the scales along 
the lateral line are more than 97 as in O. ventralis, O. attenuatus, 
and O. incomptus; in O. compressus they are 97 or less. In O. 
ceroni the upper labials are not in contact with the orbit; the same 
condition obtains in O. ventralis and in O. attenuatus, but in O. 
compressus the upper labials are in contact with the orbit; head 
scale characters are unknown in O. incomptus. The frontonasal is 
divided into an anterior and a posterior frontonasal in O. ceroni 
and in O. compressus; this scale is single in O. ventralis and O. 
attenuatus. A distinct mid-dorsal stripe is present in O. ceroni, O. 
compressus, and O. attenuatus; this stripe is absent or very indis- 
tinct in O. ventralis and O. incomptus. The area below the lateral 
fold lacks dark pigment in O. ceroni, O. ventralis, and O. incomptus; 
in O. attenuatus and O. compressus stripes or other dark markings 
occur below the lateral fold. 

Relationships. McConkey (1955) lists an Ophisaurus specimen 
(AMNH 15473) that was rendered almost useless for study by the 
blows of a machete. This specimen, from Laguna de los Cocos, 
Veracruz, was tentatively assigned to the new species O. incomptus. 
McConkey states (referring to AMNH 15473) "I should not be 
surprised if it eventually were shown to be still another distinct 
form. It is recorded from sand dunes, a habitat completely differ- 
ent from the deciduous forest area in which the Valles specimen 
was taken. My previous work with glass lizards in southeastern 
United States indicates that in the genus Ophisaurus habitat differ- 
ences are closely linked with morphological ones". In another 



314 Quarterly Journal of the Florida Academy of Sciences 

publication McConkey (1954) again refers to AMNH 15473. He 
states "It resembles attenuatus in that a mid-dorsal stripe is present, 
but is like ventralis in that the white markings form stripes run- 
ning along the edges of the scales, and in that there is no dark 
pigment below the lateral fold". These characters ally AMNH 
15473 to O. ceroni rather than to O. incomptus. 

Unfortunately, the battered condition of AMNH 15473 pre- 
cludes its definite specific allocation. Nevertheless, based on its 
morphological and habitat similarities to O. ceroni, I suggest 
AMNH 15473 should be tentatively referred to this species rather 
than to O. incomptus. 

The relationships of O. ceroni to the other New World species of 
Ophisaurus remain unclear. This is because O. ceroni has at least 
one character found diagnostic in the definition of each of the 
named forms. The rarity of Ophisaurus specimens from Mexico is 
perplexing. It is hoped that additional collecting, especially from 
coastal areas of eastern Mexico will further elucidate our knowledge 
of this interesting group of lizards. 

Our knowledge of Mexican Ophisaurus indeed remains frag- 
mentary, yet, in summary, it now appears that there are two valid 
species of glass lizard from the Neotropics of eastern Mexico. One 
of there, Ophisaurus incomptus, inhabits tropical deciduous forest; 
the other, Ophisaurus ceroni, inhabits coastal dune-scrub. 

Acknowledgments 

The trip to Veracruz in August, 1964, was made possible by 
Illinois State University Grant 64-2. Donna Rae Holman made 
the drawings. 

Literature Cited 

Auffenberg, W. A. 1955. Glass lizards (Ophisaurus) in the Pleistocene and 
Pliocene of Florida. Herpetologica, vol. 11, pp. 133-136. 

Cope, E. D. 1900. The crocodilians lizards and snakes of North America. 
Ann. Report United States Nat. Mus. for 1898, pt. II, pp. 153-1294, 
figs. 1-347, pis. 1-36. 

Etheridge, R. 1960. The slender glass lizard, Ophisaurus attenuatus, from 
the Pleistocene (Illinoian Glacial) of Oklahoma. Copeia, 1960, no. 1, 
pp. 46-47. 

. 1961. Late Cenozoic glass lizards (Ophisaurus) from the southern 

Great Plains. Herpetologica, vol. 17, pp. 179-186. 



Holm an: Neio Glass Lizard 315 

Holman, J. A. The Pleistocene herpetofauna of Saber-tooth Cave, Citrus 
County, Florida. Copeia, 1958, no. 4, pp. 276-280. 

McConkey, E. H. 1954. A systematic study of North American lizards of 
the genus Ophisaurus. American Midland Nat., vol. 51, no. 1, pp. 133- 
171. 

. 1955. A new lizard of the genus Ophisaurus from Mexico. Natural 

History Misc., Chicago Acad. Sci., no. 145, pp. 1-2. 

Smith, H. M., and E. H. Taylor. 1950. An annotated checklist and key to 
the reptiles of Mexico exclusive of the snakes. Bull. U. S. Nat. Mus., 
199, i-vi and pp. 1-253. 

Weigel, R. W. 1962. Fossil vertebrates of Vero, Florida. Florida Geol. 
Surv. Sp. Publ. 10, pp. 1-59, i-vii. 

Yarrow, H. C. 1884. Check list of North American reptilia and batrachia. 
Bull. United States Nat. Mus. 24, pp. 1-249. 

Department of Biological Sciences, Illinois State University, 
Normal, Illinois. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



SUBSPECIATION IN SPHAERODACTYLUS COPEI 
Albert Schwartz and Richard Thomas 

In 1867, Steindachner described a new species of gecko, Sphae- 
rodactylus copei from "Siidamerika, ohne nahere Angabe des Fun- 
dortes". The name was later applied to the large geckoes of south- 
western Hispaniola by Barbour (1921:259). Garman (1887) had 
described the same species as S. picturatus, a name which is pres- 
ently in the synonymy of copei Steindachner. Finally, one of us 
(Schwartz, 1961), in a review of the scaber group of the genus 
Sphaerodactylus, regarded copei as racially related to the Bahaman 
S. anthracinus. 

Through the courtesy of Dr. Ernest E. Williams, Museum of 
Comparative Zoology (MCZ), we have been able to examine not 
only the syntypes of S. picturatus Garman, but also a large lot of 
freshly collected material of copei from the southwestern peninsula 
of Haiti. In addition, we have been able to amass significant col- 
lections from the area ourselves. When all this material is as- 
sembled, it is quite obvious that within the population which is 
presently known as S. a. copei occur several very distinctive sub- 
species; an analysis of this geographic variation is the purpose of 
the present paper. We wish to thank not only Dr. Williams for 
his help in the present endeavor, but also our fellow collectors 
in southwestern Haiti, Miss Patricia A. Heinlein, and Messrs. Ron- 
ald F. Klinikowski, David C. Leber, and Dennis R. Paulson. The 
illustrations are the work of Mr. Klinikowski, and he has our sin- 
cere thanks for his efforts on our behalf. Paratypes of one new 
form have been deposited in the Carnegie Museum (CM), the 
American Museum of Natural History (AMNH), the Museum of 
Natural History, University of Kansas (KU), the Museum of Zo- 
ology, University of Michigan (UMMZ), and the United States 
National Museum (USNM). 

We feel that Schwartz's action in combining S. copei and S. 
anthracinus was premature. Not having seen live specimens of 
either species, he was completely unaware of the vivid coloration 
and pattern of copei. Although S. anthracinus is still unknown to 
us in life, and there seems to be no adequate color description of 
the species, we feel that it is a preferable course at present to once 
again separate S. anthracinus and S. copei, at least until there is 



Schwartz and Thomas: Subspeciation in Sphaerodactylus 317 

definite information as to the coloration of the former. We 
acknowledge, however, that there is a community of character- 
istics between the two species — the dark collar with four pale 
ocelli, pale head striping on a dark ground, heavily vermiculate 
heads in "adult" males, escutcheon shape — which ally these two 
species to one another more closely than to any other described 
Bahaman-Hispaniolan forms. With the description of several new 
races of S. copei from Hispaniola, it seems preferable to separate 
the two species. It is possible that, if anthracinus is maintained in 
the same species as copei, the former may not have differentiated at 
the same level as the Hispaniolan races of the latter; this is an added 
reason for the separation of the two species. 

S. copei was described from the collections made by the per- 
sonnel of the Austrian frigate Novara in its journey around the 
world. Gans (1955) has shown that the Novara did not touch any 
West Indian island, going directly from Madeira to Rio de Janeiro. 
Thus the origin of the type specimen of copei is shrouded in mys- 
tery. Karl Scherzer, one of the expedition's members, left the 
vessel at Valparaiso, journeyed alone through South America, vis- 
ited the island of St. Thomas, where he received a gift of zoolog- 
ical material from A. Riise, and rejoined the expedition at Gibral- 
tar. It is possible, after the manner of the times, that Riise had 
received through exchange a single gecko from Haiti, which in 
turn he gave (without locality data) to Scherzer, and which in 
turn became part of the Novara collections. 

Another possibility is that the name S. copei has been mis- 
applied to any Hispaniolan gecko; it may indeed be a South Amer- 
ican species (although this is extremely unlikely) or may be from 
elsewhere in the West Indies. Without further information (and 
the description is extremely detailed but uselessly so insofar as 
determination of the species is concerned) it is pointless to specu- 
late further. 

Assuming that S. copei is indeed Hispaniolan, and knowing that 
the species is geographically variable there, it is necessary to re- 
strict the name to one of the four distinct Hispaniolan populations. 
No pertinent scale counts are given in the description. The color- 
ation is described as "upperside of head behind the eyes brownish 
with gray flecks, the forehead and snout unicolor brownish; back 
evenly flecked with brown and gray; venter bright brownish gray 
with brownish scale edges." The specimen is obviously a plain 



•318 Quarterly Journal of the Florida Academy of Sciences 

male; it may have had a partially vermiculate head in life. The 
description of coloration and pattern is not distinctive, and the 
specimen may have come from any of the Haitian seaport towns 
along the southern peninsula, namely Port-au-Prince, Miragoane, 
Jeremie, Les Cayes, or Jacmel, at or near all of which the species 
is now known to occur. Since the name has priority and has been 
used by various authors, it must be conserved. Purely because of 
convenience and the vague possibility that the type may have come 
from the vicinity of the Haitian capital, we restrict the type locality 
of S. copei Steindachner to the vicinity of Port-au-Prince, Dept. de 
TOuest, Haiti. 

Sphaerodactylus copei Steindachner 

Diagnosis. A sphaerodactyl with a middorsal zone of granules 
and large keeled and imbricate dorsal scales. Ventrals smooth and 
round; chest scales smooth. Gulars smooth. Internasals to 3 
(mode 1), upper labials 3. Strong sexual dichromatism: preserved 
adult males dull grayish and without pattern or with indications of 
a salt-and-pepper pattern, although in life the dorsum is blue-gray, 
brown, or greenish, with scattered rusty and/or purplish-brown 
scales, head either yellow-green or vivid rusty-orange, in one race 
developing additionally a heavily black reticulate cephalic over- 
lying pattern; female dorsal ground color varying from apparently 
pale grayish to blue-gray or dark brown, with one to four transverse 
body bands (obscure in some races) with gray scales within the 
bands, a black nuchal collar with normally four white ocelli, and 
a complex head pattern of pale (greenish or rich brown) lines on 



Plate 1. Hispaniolan geckos. Fig. 1. Sphaerodactylus copei copei, adult 
female, ASFS X2371, from Diquini, Dept. de l'Ouest, Haiti; snout-vent length 
36 mm (top row, left). 

Fig. 2. Sphaerodactylus copei enochrus, adult female, type, MCZ 65128, 
from Marbial, 21 km NE Jacmel, Dept. de l'Ouest, Haiti; snout-vent length 
38 mm (top row, center). 

Fig. 3. Sphaerodactylus copei picturatus, adult female, MCZ 69971, 
from Place Negre, near Jeremie, Dept. du Sud, Haiti; snout-vent length 35 
mm (top row, right). 

Fig. 4. Sphaerodactylus copei cataplexis, adult female, type, MCZ 77161, 
from Camp Perrin, Dept. du Sud, Haiti; snout-vent length 39 mm (bottom 
row, left). 

Fig. 5. Sphaerodactylus copei cataplexis, adult male, AMNH 92805, from 
Camp Perrin, showing heavily vermiculate head which occurs in some male 
cataplexis; snout-vent length 37 mm (bottom row, right). 



Schwartz and Thomas: Subspeciation in Sphaerodactylus 319 








320 Quarterly Journal of the Florida Academy of Sciences 

a drab (blue-gray or gray) background. Dorsal scales 14 to 22 be- 
tween axilla and groin; ventral scales 24 to 32 between axilla and 
groin; 39 to 52 scales around midbody; 10 to 17 fourth toe lamel- 
lae; escutcheon 4 to 8 X 19 to 30; size large, males to 41 mm in 
snout-vent length, females to 40 mm. 

1. Sphaerodactylus copei copei Steindachner 

Diagnosis. A subspecies of S. copei characterized by high num- 
ber of dorsal scales between axilla and groin, high number of 
scales around midbody; males pale gray-blue dorsally, dotted with 
rust and dark purplish-brown, head pale yellow-green with a faint 
rust head pattern persisting into adulthood, throat and tail yellow, 
ventral ground color purple, tail blue-gray basally; females usually 
with a black collar with four white ocelli, a more or less complete 
rich brown U enclosing a prominent and contrasting gray occipital 
spot, two complete pale nuchal bands enclosing between them- 
selves a dark nuchal band, these three bands anterior to the black 
collar, dorsal ground color blue-gray, marbled with dark purple 
and brown, two transverse body bands (enclosing gray blotches 
within themselves) rather obsolete in fully adult females, tails yel- 
low with four pale bands bordered in black, the posteriormost 
three white, the most proximal dull yellowish-gray. 

Distribution. Known only from Ca Ira, Diquini, and "south- 
west of Port-au-Prince", along the southern shore of the Golfe de 
la Gonave at the base of the Tiburon Peninsula. 

Discussion. Since females show the head pattern more clear- 
ly than do males, and since the body pattern of females is more 
complex than that of males, it is pertinent to describe the females 
of S. c. copei in detail. The head pattern consists of a series of rich 
brown (pale in preservation) markings on a blue-gray ground color, 
as follows: a median interorbital line stopping abruptly behind the 
eyes and followed by a transverse short pale bar (which may be 
reduced to two small dots), followed by the prominent ovoid 
occipital gray spot; a pair of canthal lines which proceed to the 
anterior border of the eye; a U-shaped figure from the posterior 
border of the eyes enclosing the occipital spot, and joined anterior- 
ly to a vertical postocular stripe which proceeds from the supra- 
labials to meet the anterior end of the U-shaped figure just behind 
the eye; a pair of transverse nuchal bands enclosing between them 



Schwartz and Thomas: Sabspeciation in Sphaerodactylus 321 

a dark transverse nuchal band, these three bands just anterior to 
the black collar, the anteriormost pale nuchal line joined at times 
to the U-shaped figure, which in turn may be slightly fragmented 
posteriorly, but is always more or less complete; a black collar en- 
closing four white ocelli (Fig. 1); dorsum blue-gray, mottled ("salt- 
and-pepper") with dark purple and brown; at most two transverse 
body bands between the fore- and hindlimbs, usually only the more 
posterior persisting (if either does), the typical adult female pat- 
tern thus consisting merely of a variegated dorsum with a single 
transverse darker band with some gray blotching enclosed within, 
even this single band disappearing or obliterated in some adult 
females, entire ventral surface flesh. Males pale gray-blue dor- 
sally, spotted with rust and dark purplish-brown; head pale yellow- 
green with remnants of the female head pattern rather indistinct 
rust; tail very pale yellow, throat yellow, flecked with rust; ventral 
ground color purple, tail blue-gray basally. 

Although there is only a single male presently at hand, Schwartz 
(1961, p. 20) commented that male copei had persistent remnants 
of the female head pattern; all specimens examined by him were 
from Diquini, and are thus assignable to the nominate race. Ap- 
parently male c. copei are characterized by the persistence of the 
female head pattern into adulthood. A single juvenile does not 
show the body pattern appreciably more distinct than do adult 
females. The intensity of the transverse body bands in the females 
is somewhat variable, with MCZ 64921 showing these bands most 
prominently. Even in this specimen the clarity of the bands is 
much obliterated by dark scales in the interband spaces. In most 
females the bands are absent or only very faintly discernible. The 
black collar is present in all females except one young individual 
(ASFS X2370) which appears to lack it; in life this lizard had only 
two cream ocelli, outlined in brown, on the shoulders. 

In scalation, S. c. copei (twelve specimens) has between 19 and 
22 dorsal scales (mean 20.6) between axilla and groin, between 24 
and 32 ventral scales (mean 27.3) between axilla and groin, be- 
tween 42 and 51 (mean 46.2) scales around midbody, fourth toe 
lamellae from 12 to 16 (mean 14.8, mode 15), or 1 internasal 
(mode 1), and the single male has the escutcheon 5 X 19. The 
male has a snout-vent length of 35 mm, the largest female a snout- 
vent length of 40 mm. 

Specimens examined. Haiti, Dept. de YOuest: MCZ 64921, 



G22 Quarterly Journal of the Florida Academy of Sciences 

Qa Ira; MCZ 65108-10, 9362-63, ASFS X2369-74, Diquini; MCZ 
51255-57, southwest of Port-au-Prince. 

2. Sphaerodactylus copei enochrus, new subspecies 

Type. MCZ 65128, an adult female, from Marbial, 21 kilo- 
meters northeast of Jacmel, Dept. de l'Ouest, Haiti, collected by 
L. Whiteman, 21-22 April 1961. 

Paratypes. All from Dept. de l'Ouest, Haiti. MCZ 65118-27, 
same data as type; MCZ 65137, Jacmel, L. Whiteman, 19 April 1961; 
MCZ 65134-36, Meyer, 8 km E Jacmel, L. Whiteman, 25 April 1961; 
MCZ 65129-33, Bascap Rouge, 10 km NE Jacmel, L. Whiteman, 
24 April 1961. 

Diagnosis. A subspecies of S. copei characterized by low num- 
ber of dorsal scales between axilla and groin, low number of scales 
around midbody, females with pale dorsal ground color, black collar 
and ocelli obliterated, no U-shaped cephalic figure, a pair of pale 
dots on each side of the occipital spot, first pale nuchal band re- 
duced to a transverse series of dots, second pale nuchal line ob- 
scure, transverse body bands obliterated by heavy dark salt-and- 
pepper mottling on pale ground; male coloration in life unknown 
but preserved specimens uniform tan with no dorsal mottling and 
no remnants of female head pattern. 

Description of type. An adult female with the following meas- 
urement and counts: snout-vent length 38 mm; tail broken; dorsals 
between axilla and groin 16; ventrals between axilla and groin 28; 
midbody scales 43; fourth toe lamellae 16; internasal 1. Dorsal 
ground color (in preservation) pale gray, head somewhat brownish, 
but nonetheless pale; head pattern consisting of a pale interorbital 
line, a pair of pale canthal lines, a pair of postocular lines, ending 
shortly posterior to the eyes, with an anteroventral extension to 
the supralabials; a tiny pale dot at the posterior end of the median 
unpaired interorbital line; an occipital spot with a pair of slightly 
smaller pale spots on either side, followed by a second, more or 
less transversely aligned row of spots, somewhat fainter than the 
proceeding row and representing the first pale transverse nuchal 
band; collar dull gray, with four white ocelli, both collar and ocelli 
much fragmented and inconspicuous by virtue of the dark mottling 
on the remainder of the dorsum (Fig. 2); a single transverse body 
band barely discernible behind the forelimbs; all limbs heavily 



Schwartz and Thomas: Subspeciation in Sphaerodactylus 323 

mottled with dark gray on a pale gray ground; venter immaculate 
pale; cheeks and neck gray with some spotting. 

Variation. Seventeen paratypes show the following scale 
counts: dorsals axilla to groin, 15-18 (mean 15.7); ventrals axilla to 
groin, 25-31 (mean 27.4); midbody scales, 40-45 (mean 42.2); fourth 
toe lamellae, 14-17 (mean 15.4; mode 15); internasals, 0-1 (mode 1); 
escutcheon, 6-8 X 21-29. The largest male has a snout-vent length 
of 39 mm, the largest female 38 mm. 

The seven paratypic males can be easily dismissed; they are 
presently dull uniform tan with slightly grayer heads. There are 
no pattern elements on the heads and the venters are immaculate 
pale. The color of these lizards in life remains unknown; we have 
not seen live individuals of this subspecies. 

Six paratypic females are like the type in pale dorsal ground 
color, slightly paler heads, heavily mottled dorsa with both collar 
and transverse body bands much fragmented and obliterated. In 
head pattern they are like the type as well. Several females have 
the row of nuchal dots (those which represent the first pale nuchal 
band in c. copei) more regular and prominent than the type, form- 
ing a distinct transverse row of nuchal dots. There is no other 
significant cephalic pattern variation in the female paratypes. 

Six juvenile and subadult geckoes vary from the smallest (snout- 
vent length 17 mm) which has a black nuchal band without ocelli, 
a black band across the shoulders, and three black bands between 
the fore- and hindlimbs, all separated from each other by pale 
(buffy in preservation) transverse bands. The head vaguely shows 
the adult female pattern of rows of transverse dots. The five re- 
maining subadult specimens show the female head pattern in all 
its detail, and a black collar with four ocelli and four paler body 
bands, outlined with black. These body bands obviously disinte- 
grate with increasing size, and one of the subadult specimens 
(snout-vent length 23 mm) already shows an almost typical dorsal 
female pattern. The juvenile tails are vividly banded with pale 
and dark bands, a condition which persists into the adult females 
to some degree. 

Comparisons. The two races copei and enochrus are easily 
differentiate, both on the basis of scalation and pattern; likely 
coloration is also significant, although we have no color data on 
enochrus. The two races are separable on the basis of dorsal scales 
(15-18 in enochrus, 19-22 in copei). Despite the great overlap in 



324 Quarterly Journal of the Florida Academy of Sciences 

midbody counts (40-45 in enochrus, 42-51 in copei), the lower mean 
of enochrus (42.2) is significantly different from that of copei (46.2). 
Males of the two races can presumably be distinguished by the ab- 
sence of any trace of female head pattern in male enochrus, this 
pattern persisting into adult male copei. The females of the two 
subspecies are easily distinguished by the head pattern; the U- 
shaped head figure, two pale nuchal bands enclosing a darker band, 
the black collar with ocelli of copei contrast strongly with the dot- 
ted head, dotted anterior nuchal "band", and poorly differentiated 
or obscured collar and ocelli of enochrus. 

3. Sphaerodactylus copei picturatus Garman 

Diagnosis. A subspecies of S. copei characterized by low num- 
ber of dorsal scales between axilla and groin, high number of scales 
around midbody; males (preserved) tan, occasionally with faint 
remnants of female head pattern, dorsum dotted with dark scales 
or unicolor; females with very dark heads and dark dorsal ground 
color, head pattern of three longitudinal lines (described in detail 
below), a pair of pale nuchal bands enclosing between them a 
slightly darker area, black collar with four ocelli complete and dis- 
tinct, usually two to four prominent transverse body bands each 
including gray confluent "ocelli". 

Distribution. Known only from the vicinity of Jeremie and 
apparently Grande Cayemite, Haiti. 

Discussion. Garman (1887, pp. 19-20) described Sphaerodac- 
tylus picturatus on the basis of four syntypes from "western Haiti"; 
the specimens (MCZ 3341-42) were collected by Dr. D. F. Wein- 
land at Grande Anse River (= Riviere de la Grande Anse) which 
lies just to the south of Jeremie and has its mouth immediately to 
the southeast of that city. We have examined the syntypes and 
find them to be somewhat faded, but nonetheless identical to an 
excellent series of specimens from the Jeremie area. The syntypes 
include one unicolor male, two adult females, and a subadult indi- 
vidual. 

Garman's description of the head pattern of the female syntype 
may well be quoted; our own terms for the various head pattern 
elements are included in parentheses within the Garman descrip- 
tion: "The head is marked with white in a narrow streak (canthal 
line) on each side from the rostral on the canthus and over the 



Schwartz and Thomas: Subspeciation in Sphaerodactylus 325 

supraorbitals to the back of the head, in a median streak (inter- 
orbital line) on the forehead, a rounded spot above ear, another 
on the occiput and an oblique streak behind each ear upward to 
the back of the neck." This pleasantly detailed description agrees 
with the head pattern of the fifteen females at hand. The occipi- 
tal spot is prominent and the postorbital lines converge toward it 
on either side, extending posterior to it and often joining the first 
nuchal pale band, thereby forming a large V which encloses the 
occipital spot. No specimen has a U-shaped cephalic figure; the 
posterior tip of the V in picturatus is either incomplete or formed 
by the junction of the two postorbital lines with one another or by 
the junction of these lines with the first pale nuchal band (Fig. 3). 
One female (MCZ 69901) has the dark head ground color spotted 
with pale in a rather irregular fashion, these pale spots represent- 
ing the more orthodox pale head lines which have been reduced 
to spots. The collar is bold and contains usually four white ocelli, 
although one female (MCZ 69901) has but a single white ocellus 
surrounded by an irregular ring of black on the left side of the 
neck. The body bands number from two to four and those which 
are present are usually bold and distinct against the brown ground 
color, and have gray ocelli, either separate or fused, enclosed with- 
in them. The tail is banded with about four or five pale (white) 
bands, each bordered anteriorly and posteriorly by black. The 
limbs are all heavily mottled with dark brown. 

Four juveniles show the adult female head pattern, and have 
the dorsum patterned as described for juvenile S. c. enochrus — 
a series of black transverse body bands separated by buffy bands, 
and a conspicuously black and white ringed tail. 

Five males (including one syntype) are divided between two 
which show no dorsal pattern at all and three which have the tan 
back dotted with darker brownish scales; two of these latter also 
show faint indications of the female lined head pattern and the 
third shows it even more faintly. 

In scalation, fifteen specimens of S. c. picturatus have between 
14 and 18 dorsal scales (mean 16.5) between axilla and groin, be- 
tween 24 and 31 ventral scales (mean 27.8) between axilla and 
groin, between 42 and 53 (mean 46.5) scales around midbody, 
fourth toe lamellae from 13 to 16 (mean 14.0, mode 14), 1 to 3 
internasals (mode 1), and the escutcheon measures 4 to 6 X 20 



326 Quarterly Journal of the Florida Academy of Sciences 

to 24 scales. The largest specimens of each sex have snout-vent 
lengths of 38 mm. 

Comparisons. Handicapped in that we have not seen living 
specimens of picturatus, we are unable to make comparative state- 
ments regarding coloration. However, females of the races copei, 
cnochrns, and picturatus are readily distinguishable on the basis of 
head pattern. The U-shaped figure in copei, the one or two rows 
of dots on the heads of enochrus, and the V-shaped figure and 
posteriorly progressing postorbital lines of picturatus will quickly 
distinguish females of these three races. The dorsal ground color 
of enochrus is much paler than that of picturatus, and the very 
pale heads of enochrus contrast markedly with the very dark heads 
of picturatus. No male enochrus shows any remnants of the female 
head pattern, a condition which occurs in some male picturatus. 
The distinct body bands and collar of picturatus contrast with the 
indistinct collar and body bands of enochrus. From copei, female 
picturatus can be easily distinguished by the presence of transverse 
body bands in adults; although these bands occur in some female 
copei, they are regularly much more indistinct or faint than in 
picturatus. 

Scalewise, copei and picturatus are completely separable on the 
number of dorsals in the axilla-groin distance (14-18 in picturatus, 
19-22 in copei). Picturatus can be differentiated from enochrus on 
the basis of higher number of midbody scales in the former (pic- 
turatus, 42-52, mean 46.5; enochrus, 40-45, mean 42.2). 

A single female from Grande Cayemite (MCZ 25414) appears to 
be indistinguishable in pattern from S. c. picturatus; the condition 
of the specimen precludes taking dorsal or midbody scale counts. 
We regard this single lizard as representing S. c. picturatus on 
Grande Cayemite. 

Specimens examined. Haiti, Dept. du Sud: MCZ 3341-42 (four 
specimens), Grande Anse River; MCZ 69898-904, 69906-21, Place 
Negre, nr. Jeremie; MCZ 25414, Grande Cayemite. 

4. Sphaerodactylus copei cataplexis, new subspecies 

Type. MCZ 77161, an adult female, from Camp Perrin, Dept. 
du Sud, Haiti, one of series collected by D. C. Leber, R. F. Klini- 
kowski, and natives, 22 July 1962. Original number X2765. 

Paratypes. ASFS X2639-50, X2759-64, X2766-72, UIMNH 



Schwartz and Thomas: Sabspeciation in Sphaerodactylus 327 

55606-20, UMMZ 125200-19, MCZ 62549-60, Camp Perrin, Dept. 
du Sud, Haiti, S. Rand and J. D. Lazell, 6 August 1960; ASFS 
X2515-19, Camp Perrin, Dept. du Sud, Haiti, R. F. Klinikowski, 22 
July 1962; AMNH 92800-11, CM 39445-64, KU 79828-43, USNM 
151755-74, Camp Perrin, Dept. du Sud, Haiti, natives, 24 July 1962. 

Associated specimens. All from Haiti, Dept. du Sud. ASFS 
X3138, X3314, X3550, X3711, X3858-59, X4041-43, X4633, Camp 
Perrin — all hatchlings from eggs collected at Camp Perrin; ASFS 
X3274-76, Carrefour Canon, 500'; MCZ 63239-48, ASFS X3373, 
Les Cayes; ASFS X3792-93, 4.5 mi. NW Les Cayes; ASFS X3685- 
708, Cavaillon; ASFS X3556-63, Ile-a-Vache, western end. 

Diagnosis. A subspecies of S. copei characterized by moderate 
number of dorsals between axilla and groin, moderate number of 
scales around midbody, females with dorsal ground color greenish- 
gray, speckled with rusty scales, dark head with three creamy- 
green longitudinal lines, prominent black collar with from two to 
four white ocelli, one or two creamy-green spots on each side of 
occipital spot, no dark nuchal band but anterior pale nuchal band 
either more or less complete or reduced to dots and posterior pale 
nuchal band absent, and one to three dark transverse body bands 
present and enclosing white scales; males with rusty-orange heads, 
at times with pale green markings; dorsum green anteriorly, be- 
coming dark brown anterior to sacrum, anterior body dotted with 
rusty, tails dull blueish, throat vivid orange flecked with yellow; 
some males with a dark blue to black heavy reticulum over the 
entire top of head onto neck, and over chin and throat as well. 

Description of type. An adult female with the following meas- 
urements and counts: snout-vent length 39 mm, tail 27 mm, regen- 
erated; dorsals between axilla and groin 19; ventrals between axilla 
and groin 28; midbody scales 44; fourth toe lamellae 13; internasal 
1. Dorsal ground color (in life) dull greenish-gray, including head, 
with some scattered rusty scales on the back; head pattern consist- 
ing of a pale (pale creamy-green) interorbital line, a pale canthal 
line, and a pair of short postocular lines, each of which has a dis- 
sociated pale dot at its posterior extremity, these dots followed by 
another dot at the same level as the large occipital spot, thus 
forming a transverse line of three pale dots across the occiput; 
a short transverse pale line between the posterior end of the inter- 
orbital line and the occipital spot; cheeks and sides of neck with 
three pale spots; a row of four pale spots across the neck at the 



328 Quarterly Journal of the Florida Academy of Scdznces 

level of the first pale transverse nuchal band; black collar distinct, 
with four white ocelli (Fig. 4); a single fairly distinct transverse 
body band between the limbs, made up of two white ocelli with 
their surrounding black pigment; two other body bands, one axil- 
lary and one sacral, barely discernible; ventral ground color green- 
ish-gray; center of throat clear gray, lips dull green; all limbs rather 
mottled dorsally with dark; regenerated tail irregularly marbled 
or mottled with dark brown; iris pale straw in life. 

Variation. Forty paratypes show the following scale counts: 
dorsals axilla to groin, 15-22 (mean 18.3); ventrals axilla to groin, 
24-31 (mean 26.9); midbody scales, 39-49 (mean 44.0); fourth toe 
lamellae, 10-15 (mean 13.1; mode 13); internasals 0-1 (mode 1); 
escutcheon, 4-8 X 22-30. The largest male has a snout-vent length 
of 41 mm, the largest female 40 mm. 

The very large series of paratypes of both sexes reveals very 
little variation in pattern. In seventy-six females, the coloration 
and pattern are as described for the type, with a few exceptions. 
The head pattern is quite constant; a few individuals have a com- 
plete or almost complete first pale nuchal band instead of a series 
of dots as does the type. The maximum number of body bands is 
four; those body bands which are present are distinct and contain 
white scales to give an ocellate effect within each band. The black 
collar may be reduced, and may have as few as only a single pale 
ocellus present. Occasional specimens have only the ocelli and 
lack the black collar almost entirely. On complete and original 
tails there are three distal white rings, each bordered by black, 
and separated by wider areas of brown; the tail thus has a dis- 
tinctly ringed appearance. 

Thirty-nine subadults, juveniles, and hatchlings show the juve- 
nile pattern and the transition therefrom to that of the adult fe- 
male. The head markings in the young are like those of the female, 
even showing the same variation in the condition — i.e., complete 
or nearly so, or dotted — of the first pale nuchal band. The collar 
lacks ocelli, and there are three or four dark body bands, enclosing 
on one hand between them dark brownish bands, and separated by 
pale buffy bands from one another, in an alternating pattern; the 
net result of this configuration is two wide dark bands including 
between them a brown band, these two triads separated by a pale 
buffy band. The collar is preceeded by a pair of buff bands (the 
first and second nuchal bands) which enclose between themselves 



Schwartz and Thomas: Subspeciation in Sphaerodactylus 329 

a darker nuchal band. With increasing age and size, the black 
body bands and collar develop gray to white centers (ocelli) and 
the definite juvenile banding becomes more obscure until, in adults, 
the transverse body bands either persist as such or disappear. 

Seventy-nine males show two patterns. The majority of the 
specimens in life had rusty-orange heads (occasionally with pale 
green linear female markings which are almost not apparent in 
the preserved specimens), dorsal ground color green anteriorly, 
dotted with rusty, turning to dark brown anterior to the sacrum. 
The tails were dull blueish, the throat a vivid orange (PI. 4 D 12, 
Maerz and Paul) flecked with yellow; the ventral color was blue- 
gray. The balance of the males (eighteen individuals) show some 
sort of head markings, ranging from a dark brown spotting or 
reticulum to a complete blue-black reticulum covering the entire 
head and throat on a grayish ground color above and yellow-orange 
below (Fig. 5). These males are not all large, nor are they neces- 
sarily the largest specimens of the entire lot. Interpretation of this 
head condition is difficult. It is possible that this is a breeding 
coloration, although it is highly doubtful that those large males 
which lack it were not completely adult. The dark head cannot 
be interpreted as a senile or "super male" character since the series 
of males so marked include specimens with snout-vent lengths as 
small as 30 mm, well below maximum size (41 mm) for males of 
this race. Both one small and one large individual show what we 
interpret as the intermediate condition, wherein the head has a 
dark brown reticulum rather than a blue-black one. The speci- 
mens look remarkably like extra-large size Guadeloupe S. fantasti- 
cus fantasticus. No other subspecies of S. copei is known to have 
this condition, although the number of males of each of the other 
races is very small and lack of large series may give a fallacious 
impression. It is of interest to note that S. anthracinus males may 
on occasion develope a comparable dark vermiculate head pattern 
(Schwartz, op. cit. :20). 

Comparisons. No scale counts will separate cataplexis from 
any other race. The mean for dorsals is intermediate between 
picturatus and enochrus on one hand and copei on the other; the 
mean for midbody scales falls between that of picturatus and copei 
on one hand and enochrus on the other. The mode of 13 (rather 
than 14 or 15) for fourth toe lamellae is of interest. 

Female cataplexis can be distinguished with ease from females 



330 Quarterly Journal of the Florida Academy of Sciences 

of the other races. Copei has a U-shaped head figure, picturatus a 
cephalic V, and enochrus two rows of dots on a pale head. The 
head pattern of enochrus and cataplexis are somewhat comparable; 
however, the distinct body bands of cataplexis along with its dark 
dorsal color distinguish it immediately from enochrus. A casual 
glance at the color descriptions of copei and cataplexis makes it 
at once apparent that these two forms are very different in color- 
ation, as well as pattern. Doubtless, if the coloration of enochrus 
and picturatus in life were known as well, these races would be 
seen to be equally distinct from cataplexis. 

Male cataplexis lack the salt-and-pepper condition of picturatus, 
lack the distinct head lines of male copei, and may have darkly ver- 
miculate heads, a condition which does not occur in any of the 
other three races. Again, coloration of living cataplexis and copei 
males reveals at once that these two forms are easily distinguish- 
able in life. 

A small series of eight specimens (four females, three males, 
one juvenile) from Ile-a-Vache does not appear to differ from speci- 
mens of cataplexis from Les Cayes on the adjacent mainland. 
None of the males in this lot shows the vermiculate head pattern. 
We regard these specimens as S. c. cataplexis. 

There are twelve specimens from the vicinity of Miragoane, as 
follows: MCZ 25411-13, Miragoane, Dept. du Sud, Haiti; MCZ 
66226-28, Butete, nr. Miragoane; MCZ 66229, Nan Corosse, nr. 
Miragoane; MCZ 66230, Risque, nr. Miragoane; MCZ 66231, Min- 
grette, nr. Miragoane; MCZ 66232-34, Pemel, nr. Miragoane. Al- 
though these specimens are intermediate geographically between 
the ranges of copei and picturatus they are much closer in pattern 
to cataplexis, which occurs to the southwest. The dorsal and head 
patterns of the females are similar to those of females from Camp 
Perrin; two of the males have the heads dotted with dark brown, 
a condition which is not precisely matched in any of the Camp 
Perrin series, but which may be an initial step between the orange 
head and the heavily vermiculate head in cataplexis. In scale 
counts these lizards are closer to enochrus (the range of which lies 
to the southeast of Miragoane) in number of dorsals (15-17), but 
closer to cataplexis in midbody scales (41-48). Although these 
specimens are somewhat problematical, it is preferable to call 
them S. c. cataplexis at least until living members of this Miragoane 
population can be observed. 



Schwartz and Thomas: Subspeciation in Sphaerodactylus 331 

Discussion 

Sphaerodactylus copei has been shown to be divisible into four 
very distinct races on the basis of coloration, head pattern, and 
scalation. The species is known to occur only on the Tiburon Pen- 
insula in southwestern Hispaniola, and presently is unknown from 
the affiliated Peninsula de Barahona in the Republica Dominicana. 
S. copei is thus completely restricted to the south island and does 
not occur in, or even closely approach, the Cul-de-Sac plain. In 
actuality the species appears to be absent from the higher moun- 
tains in the Massif de la Selle and Massif de la Hotte; thus the 
northern (copei and picturatus) populations are effectively isolated 
from the southern (enochrus and cataplexis) populations, except 
that apparently cataplexis has crossed the mountains in the Fond- 
des-Negres region and occurs on the north coast as well, between 
the ranges of copei and picturatus. Both picturatus and cataplexis 
have insular populations, the former on Grande Cayamite, the lat- 
ter on Ile-a-Vache, which have not differentiated from the popu- 
lations on the (more or less) adjacent mainland. 

Only two species of Sphaerodactylus are known from the Ti- 
buron Peninsula: S. copei and S. cinereus, although Cochran (1941, 
p. 112) listed a specimen of S. stejnegeri, collected by W. L. Abbott 
in "southwestern Haiti". During the period of 1917-18, Abbott did 
indeed collect extensively on the Tiburon Peninsula near Jeremie, 
Moron, and Moline, but between March 5 and 12, 1918, he col- 
lected near Etang Saumatre and Trou Caiman, (see Wetmore and 
Swales, 1931:25-26) both localities in the Cul-de-Sac whence S. 
stejnegeri is known. It is thus not unlikely that S. stejnegeri, typi- 
cally a xeric region lizard, does not occur on the Tiburon Peninsula. 
Neither of these species (and in fact, no Sphaerodactylus) has been 
reported from the adjacent He de la Gonave, an island whose fauna 
shows some distinct relationships with that of the Tiburon Penin- 
sula. Apparently S. copei has evolved on the western two thirds 
of the south island, has differentiated both on the north and south 
sides of the central mountain massifs, and has been able to cross 
these ranges at least in the relatively low Fond-des-Negres area. 
It would not be surprising if S. copei were to be taken in the region 
of the Vallee de Trouin; specimens from this area may well be 
found to be intergrades between the races copei and enochrus. 

At least in our experience, S. copei is distinctly not a gecko of 
xeric situations. At Camp Perrin they were extremely abundant, 



332 Quarterly Journal of the Florida Academy of Sciences 

occurring under almost any ground cover, and in piles of rocks 
and in old stone walls. At Diquini a small series was taken under 
rocks on the floor of the well-known cave at that locality. Camp 
Perrin lies at an elevation of about 1000 feet and, although not 
extremely wet, is certainly mesic and presumably was once for- 
ested. The Diquini area likewise is moderately mesic, with low- 
land cut-over forest adjacent to the cave. The altitudinal distribu- 
tion of S. copei varies from sea level (Ca Ira, Ile-a-Vache, Les 
Cayes) to 1000 feet (Camp Perrin); it is possible that some of the 
unlocatable Miragoane localities may be slightly higher than Camp 
Perrin. 

Literature Cited 

Barbour, Thomas. 1921. Sphaerodacttjhis. Mem. Mus. Comp. Zool., vol. 
47, no. 3, pp. 217-278, 26 pis. 

Cochran, Doris M. 1941. The herpetology of Hispaniola. Bull. U. S. Nat. 
Mus., no. 177, pp. 1-398, 12 pis., 120 figs. 

Gans, Carl. 1955. Localities of the herpetological collections made during 
the "Novara Reise". Ann. Carnegie Mus., vol. 33, pp. 275-285. 

Garman, Samuel. 1887. On West Indian Geckonidae and Anguidae. Bull. 
Essex Inst., vol. 19, pp. 17-24. 

Maerz, A., and Rea Paul. 1950. A dictionary of color. McGraw-Hill Book 

Co., New York., vii + 208 pp., 56 pis. 

Schwartz, Albert. 1961. A review of the geckoes of the Sphaerodactylus 
scaber group of Cuba. Herpetologica, vol. 17, no. 1, pp. 19-26, figs. 
1-10. 

Steindachner, Franz. 1867. Reise der osterreichischen Fregatte Novara 
um die Erde. Zoologische Theil, erster Band. Reptilien, pp. 18-19 (de- 
scription of S. copei only). 

Wetmore, Alexander, and Bradshaw H. Swales. 1931. The birds of 
Haiti and the Dominican Republic. Bull. U. S. Nat. Mus., no. 155, pp. 
1-483, 26 pis., 2 figs., map. 

10000 S. W. 84th Street, Miami, Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



ISOLATING MECHANISMS IN SNAKES 
Wilfred T. Neill 

Mechanisms bringing about reproductive isolation have often 
been investigated in frogs (Mecham, 1961), but not in other herpe- 
tozoan groups. In the current state of knowledge it would be im- 
possible to generalize about isolating mechanisms in the sala- 
manders, turtles, or lizards. The present paper summarizes the 
mechanisms that are thought to exist among snakes. The sum- 
mary is based largely upon personal observation in field and lab- 
oratory, supplemented by a review of the literature. The biblio- 
graphic references are not intended to be exhaustive. Where pos- 
sible, common names as well as scientific ones are employed, to 
render the account more meaningful to the non-herpetologist. 

The study is concerned primarily with mechanisms that operate 
at the species level; that bring about the reproductive isolation of 
two closely related, usually congeneric, species. 

Account of Mechanisms 

1. The most obvious form of reproductive isolation is geo- 
graphic. Two species, however compatible otherwise, will not 
interbreed in nature if their respective ranges are not in contact 

Congeners are often allopatric in snakes, but this need not be 
the case. For example, the snake fauna of Alachua County, Flor- 
ida, includes five monospecific genera (Cemophora, Drymarchon, 
Liodytes, Seminatrix, Stilosoma); two genera each with two species 
in the county, and no additional species in other areas (Farancia, 
Virginia); seven genera each with only one species in the county, 
but with one or more additional species elsewhere (Agkistrodon, 
Diadophis, Micrurus, Opheodrys, Pituophis, Sistrurus, Urotheca); 
eight genera each with two species in the county, and with one or 
more additional species elsewhere (Coluber, Crotalus, Elaphe, Het- 
erodon, Lampropeltis, Storeria, Tantilla, Thamnophis); and one 
genus with five species in the county and additional species else- 
where (Natrix). 

Range maps for North American snakes (Wright and Wright, 
1957) show both allopatry and sympatry among congeners. 

2. Two congeneric snakes may be sympatric, but confined to 
different habitats and therefore unlikely to interbreed. 



334 Quarterly Journal of the Florida Academy of Sciences 

Thus, in Marion County, Florida, the mud snake (Farancia 
abacura) inhabits acidic ponds, bogs, and swamp streams, while 
the rainbow snake (F. erytro gramma) inhabits basic (calcareous) 
spring runs (Neill, 1964, pp. 272-274, fig. 5). In Richmond County, 
Georgia, the cottonmouth (Agkistrodon piscivoms) is aquatic and 
riparian, while the copperhead (A. contortrix) is terrestrial on higher 
ground bordering stream valleys. In the Florida Panhandle, the 
glossy watersnake (Natrix rigida rigida) is confined to ponds, while 
the queen snake (N. septemvittata septemvittata), a near ally, is 
confined to streams. Throughout their area of sympatry in Georgia 
and Florida, the coach whip (Coluber flagellum) inhabits the drier 
situations, the blacksnake (C. constrictor) the damper ones. Many 
other examples could be listed. 

But it is seldom that two sympatric, congeneric snakes are so 
divergent in habitat that they cannot occur together in some inter- 
mediate situation. I have collected the mud snake and rainbow 
snake together in large, spring-fed streams with swampy borders 
and tributaries; the cottonmouth and copperhead under the same 
bridge; the coachwhip and blacksnake in wiregrass flatwoods 
(Pinus-Aristida community), which vary from very wet to very dry. 
In some cases, two closely related species occur very commonly 
in the same habitat. Thus the red-bellied watersnake (Natrix ery- 
throgaster erythrogaster) and the banded watersnake (N. fasciata 
fasciata) are found together in and beside lakes, ponds, swamps, 
and streams in the Atlantic Coastal Plain of Georgia. 

Carr (1940) described the respective habitats of Florida snakes. 

3. Two sympatric, congeneric snakes may occupy the same 
habitat, yet fail to interbreed because they seek different micro- 
habitats during the mating season. 

Thus, as noted, the red-bellied watersnake and the banded 
watersnake occur together in aquatic and riparian situations of the 
Atlantic Coastal Plain in Georgia; but in that region the former 
always mates in shallow water, while the latter always leaves the 
water to mate. Mating pairs of the banded watersnake are often 
found as much as 30 to 40 feet back from the water, where indi- 
viduals of the species are not otherwise to be expected. 

A slightly different situation involves the brown watersnake 
(Natrix taxispilota taxispilota) . In the Atlantic Coastal Plain of 
Georgia it is confined to larger streams and lakes, where it is as- 
sociated with the red-bellied and banded watersnakes. All three 



Neill: Isolating Mechanisms in Snakes 335 

species commonly bask on tree limbs overhanging the water; but 
the brown watersnake, unlike the other two, also mates in this 
arboreal milieu. Arboreal mating has been reported for the brown 
watersnake in Florida as well (Carr, 1940, p. 89). 

Separation by mating microhabitat is fairly common among 
snakes, but has not previously been discussed in the literature. 

4. Two sympatric, congeneric snakes, occupying the same hab- 
itat, may be prevented from interbreeding by the timing of their 
respective reproductive cycles. 

In Richmond County, Georgia, the blacksnake is the first snake 
to emerge from hibernation in the spring, appearing as early as 
latter February and mating soon after. But in the same region, 
the coachwhip is the last species to emerge, rarely appearing be- 
fore April, by which time the blacksnake has finished mating. 

The isolation of sympatric, congeneric snakes, through inter- 
specific differences in mating season, has been observed for several 
genera of the Southeast, but has not been discussed in the litera- 
ture. Fox (1954) investigated the proximate causes of interspecific 
differences in the timing of the reproductive cycle among certain 
California gartersnakes (Thamnophis). Neill (1962) summarized 
observations on the ultimate causes of the ophidian reproductive 
cycle. 

5. Certain aspects of ophidian courtship behavior function to 
discourage interspecific mating. 

Many accounts of supposed courtship in snakes (Davis, 1936) 
are now known to relate actually to the "combat dance" of two 
males (Shaw, 1951); but some snakes do perform fairly elaborate 
courtship antics (Noble, 1937). The true courtship has been studied 
in only a minority of snake species, and further work may reveal 
some exceptions to the following generalizations. 

The male snake is the more active in courtship, seeking out 
the female. The courting male is especially stimulated to follow 
the scent trail of an oestrous female of his own species (Noble, 
1937; Noble and Clausen, 1936). Commonly, several males find 
and try to mate with one female (Finneran, 1949). The female is 
apt to reject the first few advances; she dashes away, the males 
pursuing. Having overtaken the female, the courting male rubs 
his chin along her back, while repeatedly flickering his tongue in 
and out. Finally his chin is appressed to her nuchal region. Usu- 
ally the male makes very sharp twitching or jerking motions, and 



336 Quarterly Journal of the Florida Academy of Sciences 

undulant body movements that begin caudally; there is some inter- 
twining or overlapping of bodies, and the male lifts the female's 
cloacal region to permit intromission. 

In some species the undulant body movements are lacking and 
the male bites the female along the body, especially in the nuchal 
region. In the blacksnake, both sexes dash about before settling 
down to the Liebesspiel (Noble, 1937). 

Courtship may continue for days before the female is aroused 
to the point of acceptance. In one case a captive male red-bellied 
watersnake courted almost continuously for a month without arous- 
ing the female (Carr, 1940, p. 89). Repeated rejection of a male 
may often reflect the circumstance that the female has not become 
physiologically or psychologically ready for mating. However, this 
need not always be the case; a female may reject one male only 
to accept another (Munro, 1948). 

In captivity, enforced proximity with other species could en- 
courage an abnormal amount of attempted cross-mating; but on 
the other hand, debilitating or highly unnatural cage conditions 
could inhibit mating activity of any kind. In my experience, a 
healthy male snake, if well adjusted to captivity, if exposed to 
some natural fluctuations of temperature and illumination, and if 
deprived of a normal mating outlet, will attempt a cross-mating 
with a congener, or even with a distantly allied species, during the 
male's normal mating season. However, certain species are far 
more persistent in their cross-mating efforts than are others. Prob- 
ably the least discriminating is the eastern hognose snake (Hetero- 
don platyrhinos); captive males of this species will attempt to mate 
with almost any kind of snake, of either sex. 

Male gartersnakes, courting their own kind, ignored anes- 
thetized or freshly killed females (Noble, 1937); but male hognose 
snakes attempted to mate with a dead female of their own kind 
(Medsger, 1927), and a male lancehead snake (Bothrops jammed) 
attempted a necrophilous intergeneric mating (Amaral, 1932). 

A minority of snake species are provided with special anatomi- 
cal structures that function during courtship and mating. Chin 
tubercles, which hypertrophy during the mating season, may serve 
a hedonic function (Blanchard, 1931). Supracloacal tubercles serve 
a tactile function (Blanchard, 1931; Noble, 1937). The hind limb 
vestiges of certain boas (Boidae) are used by the male to scratch 



Neill: Isolating Mechanisms in Snakes 337 

or stroke the female during courtship, and also function as claspers 
(Davis, 1936). 

Sexual dimorphism in color has been reported for about a 
dozen species of snakes (Neill, 1964, p. 282), but its possible role 
in courtship is unknown. 

Of the various aspects of ophidian courtship, three closely in- 
terrelated ones seem especially important in preventing a cross- 
mating. One is the tendency for the male to follow the scent trail 
of an oestrous female of its own species. A second is the inclina- 
tion of the female to withdraw when first approached by a court- 
ing male, and to accept a male only after some pursuit. A third 
factor is the considerable length of time necessary for a courting 
male to arouse the female to the point of acceptance. Presumably, 
a courting male would not trail a female of another species with as 
great a persistence as he would a female of his own species; and 
the longer the interspecific courtship, the greater the likelihood 
that the male will be eluded by the coy female, or will undergo 
a weakening of the trailing impulse, or will be diverted onto a 
trail of his own kind. 

6. A difference in size may function in three ways to prevent 
the interbreeding of congeneric snakes. 

A scarlet kingsnake (Lampropeltis d. doliata), usually under 45 
cm. in total length, probably could not mate with a Florida king- 
snake (L. getulus floridana) even if both individuals were stimu- 
lated to attempt the cross-mating; for examples of the latter spe- 
cies commonly exceed 150 cm. in length. 

Also, a large snake, simply by virtue of its size, might well 
evoke from a smaller congener an avoidance reaction taking pre- 
cedence over reproductive urge. 

Finally, many snakes prey upon smaller individuals of other 
species; and, within certain of the more ophiophagous genera 
(e.g., Coluber, Lampropeltis, Agkistrodon), a large snake might re- 
gard a smaller congener as prey rather than as a potential mate. 

Among snakes, two sympatric, congeneric species usually differ 
in both average and maximum size, but the difference is rarely so 
great as to rule out all possibility of cross-mating. Normal matings 
commonly involve a small male and a much larger female; for in 
most snake species, the females average decidedly larger than the 
males, and reach a greater maximum size. 



338 Quarterly Journal of the Florida Academy of Sciences 

7. Interbreeding of congeners may be limited by interspecific 
differences of the reproductive organs. 

The hemipenis of the male snake is an extraordinary structure; 
it is equipped basally with hooks or enlarged spines, and more 
distally with spines, flounces, calyces, etc., depending on species 
(Dowling and Savage, 1960). The anatomy of the female's cloaca 
must be such as to permit reception of the various spines, etc. If 
the hemipenis of the male is spinous in a given species, the cloaca 
is thick-walled in the female of that species; if the hemipenis is 
smooth, the female's cloaca is thin-walled (Cope, 1900, p. 700; Pope, 
1941). The copulating male is anchored firmly by the basal spines 
or hooks of the hemipenis (Beuchelt, 1936); the more distal orna- 
mentation must serve another function. 

Hemipenial characters were found to be constant in nineteen 
species of the snake genus Calamaria, but variable in a twentieth 
(Inger and Marx, 1962). The size of the basal hooks or spines is 
a character of subspecific import in the blacksnake (Auffenberg, 
1955). Hemipenial differences often exist at the specific and ge- 
neric levels (Dowling and Savage, 1960), and at higher levels 
(Clark, 1944). 

Two Old World species of lancehead snakes (T rimer esurus), 
externally very similar, differ so markedly in hemipenial characters, 
and correlated cloacal characters of the female, that a cross-mating 
might be mechanically impossible (Pope, 1941). Such an extreme 
difference between congeners is, however, unusual; in most cases 
a male snake, attempting an interspecific mating, probably would 
not be prevented from intromission by interspecific differences in 
hemipenial ornamentation. 

A small male of the eastern hognose snake (Heterodon platy- 
rhinos) mated with a large female of the southern hognose snake 
(H. simiis); the female died three days later, possibly from cloacal 
lacerations produced by the hemipenial spines of the male (Neill, 
1951, pp. 52-53, fig. 2). 

Pope (1941, as Liophis) examined a pair of Leimadophis poe- 
cilogyms, preserved and fixed while in copula. The tips of the 
divided sulcus spermaticus were closely appressed to the openings 
of the oviducts, and the hemipenial ornamentation gripped the 
female's cloacal walls so firmly that no shifting from this position 
was possible. The circumstance suggests the probability that in- 
semination is accomplished more effectively in a normal mating, 



Neill: Isolating Mechanisms in Snakes 339 

with close hemipenial-cloacal correspondence, than in an inter- 
specific mating without such correspondence. 

8. An interspecific mating may prove fruitless, or relatively 
so as compared with a normal mating, in consequence of physio- 
logical incompatibility. By this is not meant genetic incompat- 
ibility, but the failure of the sperm to survive in an alien environ- 
ment: the reproductive tract of a female snake belonging to an- 
other species. 

When fertilization immediately or shortly follows mating, this 
factor may not be important. However, sperm storage is carried 
on by various snakes (Neill, 1962, pp. 247-248). Indeed, the females 
of all snakes may be provided with seminal receptacles (Fox and 
Dessauer, 1962, pp. 595). In one case, a single copulation sufficed 
for the production of fertile eggs in at least five successive years 
(Haines, 1940). One would suppose the longevity of sperm to be 
reduced in an alien seminal receptacle. 

Although an ectotherm, a snake, at least during its active sea- 
son, maintains a fairly constant body temperature by restricting its 
activity to times when, and places where, certain environmental 
temperatures prevail; and within this thermal activity range there 
is an eccritic (i.e., "preferred") temperature (Cowles and Bogert, 
1944; Fitch, 1956). The eccritic temperature may differ between 
congeneric snakes, by a few Centigrade degrees; and the thermal 
activity range may differ even more markedly. Reptile sperm 
cells may be adversely affected by a few degrees rise of tempera- 
ture (Cowles and Burleson, 1945). Little is known of other factors 
that might conceivably affect sperm longevity in reptiles. 

9. If an interspecific mating is accomplished, its effectiveness 
may be limited by genetic incompatibility. 

There were a few early accounts of supposed hybridization 
between snakes of remotely related genera, such as Natrix X 
Liophis or even Natrix X Vipera (Mertens, 1950). This was prior 
to the discovery of sperm storage in these reptiles. If a female 
snake, penned for a long while with a male of a different species, 
chanced to produce offspring, it was mistakenly assumed that the 
young had to be of hybrid origin. It is significant that some of 
these early accounts described the supposed hybrids as resembling 
the mother. There have been authentic instances of cross-copula 
between fairly remote relatives, e. g. Bothrops X Crotalus (Schot- 



340 Quarterly Journal of the Florida Academy of Sciences 

tier, 1950) or Pseastes X Spilotes (Mole, 1924); but these were 
without issue. 

There are a few acceptable accounts of intergeneric hybrids in 
snakes; but in each case the hybridizing genera are closely related, 
their recognition as separate entities being more a matter of 
nomenclatural convenience than of zoology. One intergeneric rat- 
tlesnake hybrid, Sistrurns X Crotalus, is known (Bailey, 1942; 
Klauber, 1956, p. 208). Possible hybrids of the mud snake and 
rainbow snake have been reported, but the two species have lately 
been placed in one genus, Farancia (Neill, 1964, p. 263). Noble 
(1937) credenced Bonnenberger's (1909) account of a Natrix X 
Thamnophis hybridization. A Thamnophis, courting its own kind, 
continued courtship when placed with a Natrix (Noble, 1937). 

Most snake hybridizations have been between congeners. Mer- 
tens (1950, 1956) has summarized much of the pertinent literature. 
Reported combinations include ratsnakes, Elaphe guttata X £• 
obsoleta; watersnakes, Natrix natrix X N. maura, N. natrix X N. 
tessellata, N. tessellata X N. maura, and N. sipedon X N. fasciata 
(Conant, 1963); gartersnakes, Thamnophis marcianus X T. radix 
(Smith, 1946); hognose snakes, Heterodon platyrhinos X H. simus 
(Edgren, 1952); vipers, Vipera aspis X V. ammodytes, and V. aspis 
X V- berus; and rattlesnakes, Crotalus adamanteus X C- horridus, 
C. unicolor X C. scutulatus, C. viridis X C- ruber, and C. viridis 
X C. scutulatus (Klauber, 1956). 

Supposed viper hybrids, Viper berus X V- ammodytes, have 
been described, but their identification as such is questioned by 
some. Two individual snakes, reported as hybrid (Thamnophis 
melanogaster X T. rufipunctatus; Bothrops cotiara X B. jararaca), 
were later interpreted in another fashion. A lancehead cross- 
copula, Bothrops jararaca X &• neuwiedi, has been observed. 

A watersnake hybridization, Natrix tessellata X N. maura, pro- 
duced only four young (Klinge, 1925). N. tessellata normally lays 
5 to 25 eggs, N. maura 4 to 20 (Mertens, 1952). A viper hybridiza- 
tion, Vipera aspis X ^ 7 - ammodytes, also produced only four young 
(Schweizer, 1941). V. aspis and V. ammodytes each normally pro- 
duces four to 18 young. In both of the above hybridizations, the 
relatively small number of offspring suggests some hybrid invia- 
bility. 

A rattlesnake hybridization, Crotalus viridis X C. ruber, pro- 
duced nine young, only one of which lived more than a year. This 



Neill: Isolating Mechanisms in Snakes 341 

one, a male, lived more than nine years, but never mated. Appar- 
ently its mechanism for sex recognition was somehow reversed, 
lor it attempted to mate with other males, and to carry on the 
male "combat dance" with females (Klauber, 1956). 

Another rattlesnake hybridization, Crotalus unicolor X C. scu- 
tulatus, produced four young, all of which were raised to maturity. 
Of these hybrids, the single female eventually mated with one or 
more of the males, producing five broods. The F 2 snakes proved 
very delicate; and autopsy revealed some of them to have anatom- 
ical defects of the stomach, heart, liver, and reproductive organs 
(Klauber, 1956). 

Yet another rattlesnake hybridization, Crotalus viridis X C. 
scutulatus, produced twelve young, some of which exhibited asym- 
metric, partially lineate patterns (Klauber, 1956). Such patterns 
appear to be strongly selected against in nature, possibly because 
they are linked with behavioral or physiological abnormalities 
(Neill, 1963, p. 209). 

The foregoing observations imply some genetic incompatibility 
among snakes at the species level, with greater incompatibility at 
higher levels. 

10. Hybrid adaptive inferiority probably exists in snakes. 

The most common snake hybridization involves Elaphe guttata 
and one of the peninsular Florida subspecies of E. obsoleta. This 
hybrid combination has been produced in captivity (Lederer, 1950; 
Mertens, 1950). Natural hybrids between the red ratsnake ("corn 
snake," E. guttata guttata) and the Everglades ratsnake (E. obsoleta 
rossalleni) have been reported (Neill, 1949, p. 10). Such hybrids 
are now known to be fairly common about certain small towns on 
or near Lake Okeechobee in southern Florida. 

Under natural conditions the Everglades ratsnake inhabits open 
sawgrass marshes, and also climbs frequently into trees and shrubs. 
It has a proportionately long tail. Its pattern is usually lineate, 
occasionally unicolor. A long tail characterizes most arboreal 
snakes; and a lineate or unicolor pattern is common among snakes 
that are arboreal, or that live in graminoids. 

In contrast, the red ratsnake, while occasionally climbing about 
tree stumps and bushes, is often subterranean, frequenting mam- 
mal burrows; it also prowls on the surface of the ground, usually 
in wooded areas. It has a proportionately short tail and a blotched 
pattern. A short tail characterizes the more secretive or subter- 



342 Quarterly Journal of the Florida Academy of Sciences 

ranean snakes, and a blotched (disruptive) pattern is usual among 
snakes that are terrestrial in wooded areas. 

Thus, the aforesaid interspecific differences, in tail length and 
pattern, are probably adaptive. The natural hybrids, all apparent- 
ly of the Fi generation, exhibit an intermediate tail length; and 
their pattern includes the blotches of Elaphe g. guttata superim- 
posed, in pallid version, upon the stripes of E. obsoleta rossalleni. 
The hybrids are therefore not ideally adapted for the natural habi- 
tat of either parent. 

It is probably significant that the Elaphe hybrids are collected 
mostly around human settlements, where the activities of man 
have reduced the number of organisms that prey upon, or compete 
with, the ratsnakes; and where unnatural environments, such as 
rodent-infested outbuildings, are frequented by both species of 
Elaphe. 

Possible Additional Mechanisms 

Two other isolating mechanisms should be listed separately 
because under special circumstances they might function to en- 
courage hybridization rather than to prevent it. 

In temperate regions many snakes overwinter communally. In 
the southern United States only small aggregations are usually 
formed; but in colder regions, hundreds of snakes may inhabit a 
single den. Thus a den in Utah yielded 930 Great Basin rattle- 
snakes (Crotalus viridis), 632 striped racers (Coluber taeniatus), 
127 blue racers (C. constrictor), and 41 individuals representing 
four other species (Woodbury, 1951). With the advent of warmer 
weather the snakes emerge; and, in many species, mating activities 
begin before the individuals have dispersed from the den. 

In the case of the aforesaid den in Utah, the tendency to aggre- 
gate would serve as an isolating mechanism as regards the rattle- 
snake; for an individual in reproductive condition would probably 
find and mate with a member of its own species near the den. With 
the racers, however, the aggregation of two congeners would favor 
hybridization if the usual isolating mechanisms chanced not to 
function. 

A second possible isolating mechanism has not been discussed 
in the literature. Both published accounts and personal observa- 
tion reveal that, among captive snakes, most attempted mismat- 
ings — interspecific, necrophilous, and homosexual — result from the 



Neill: Isolating Mechanisms in Snakes 343 

activity of a young male, probably in its first season of breeding. 
Perhaps the threshold of reproductive activity is lower in the young 
male. However, I have wondered if there might not be a process, 
roughly comparable to imprinting, whereby the male's reproduc- 
tive activity is modified by the first reproductive experience. At 
any rate, some force seems, with age, to direct the reproductive 
efforts of the male into proper channels, so that older males rarely 
attempt any but a normal mating. 

TABLE 1 

Isolating mechanisms in snakes 



I. Extrinsic 

1. Geographic Separation 

2. Habitat Separation 

3. Microhabitat Separation at Mating Time 

II. Intrinsic 

4. Timing of Reproductive Cycle 

5. Courtship 

6. Size Difference 

7. Incompatibility of Reproductive Organs 



8. Physiological Incompatibility 

9. Genetic Incompatibility 

10. Hybrid Adaptive Inferiority 



Mechanisms above the broken line are pre-mating, the others post-mating. 
Doubtful or occasionally operative mechanisms are omitted. 

Discussion 

Both geographic and habitat separation need not have evolved 
solely as isolating mechanisms, for they serve also to reduce inter- 
specific competition. 

Movement into a special microhabitat, at the time of courtship 
and mating, is probably more common than published observations 
would suggest, not only among snakes but also among other verte- 
brate groups. Many animals are commonly encountered in the 
field, yet are almost never seen engaged in reproductive activities. 
Of course, in some cases the animals may, at the time of mating, 
simply invade a microhabitat where they are exceptionally well 
concealed from predators. 



344 Quarterly Journal of the Florida Academy of Sciences 

Although snakes are most closely related to lizards, the two 
groups show no great similarity in courtship behavior. The court- 
ship of lizards has been investigated by Noble and Bradley (1933). 

Wholly sterile hybrids have not been observed in snakes. Hy- 
brids between the red ratsnake and the Everglades ratsnake ex- 
hibit marked hybrid vigor. 

Occasional hybridization, producing adaptively inferior hybrids, 
is not necessarily dysgenic, for it results in selection for greater 
isolation; and there is at least a possibility that the new genotype 
could exploit some abrupt change in the environment (Epling, 
1947). 

Summary 

Isolating mechanisms have been listed in a variety of arrange- 
ments (e.g., Dobzhansky, 1951; Mayr, 1942). The present arrange- 
ment (Table 1), based upon the snakes, is useful in that the various 
mechanisms may be thought of as operative in sequence. Thus, 
if two congeneric species are separated geographically, they can- 
not interbreed; if they are not so separated, they may yet be iso- 
lated by restriction to different habitats; if frequenting the same 
habitat, they may still be separated by invasion of different micro- 
habitats in the mating season; and so on down the list. This ar- 
rangement also permits grouping into extrinsic vs. intrinsic mech- 
anisms, and into pre-mating vs. post-mating ones. 

Literature Cited 

Amaral, A. do. 1932. Contribucao a biologia dos ophidios do Brasil. IV. 
Sobre um caso de necrophilia heterologa na jararaca (Bothrops jararaca). 
Mem. Inst. Butantan, vol. 7, pp. 93-94. 

Auffenberg, W. 1955. A reconsideration of the racer, Coluber constrictor, 
in eastern United States. Tulane Stud. Zool., vol. 2, no. 6, pp. 89-155. 

Bailey, R. M. 1942. An intergeneric hybrid rattlesnake. Amer. Nat., vol. 
76, pp. 376-385. 

Beuchelt, Hans. 1936. Bau, Funktion und Entwicklung der Begattungsor- 
gane der mannlichen Ringelnatter (Natrix natrix L.) und Kreuzotter 
(Vipera herus L.). Morph. Jahrb., vol. 78, pp. 445-516. 

Blanchard, F. N. 1931. Secondary sex characters of certain snakes. Bull. 
Antivenin Inst. Amer., vol. 4, no. 4, pp. 95-105. 

Bonnenberger, H. 1909. (Vereins-Nachrichten: Niirnberg. "Heros" E. V.) 
Bl. Aquar. Terrkde., vol. 20, pp. 827-828. 



Neill: Isolating Mechanisms in Snakes 345 

Carr, A. F., Jr. 1940. A contribution to the herpetology of Florida. Univ. 
Florida Publ., Biol. Sci. Ser., vol. 3, no. 1, pp. 1-118. 

Clark, H. 1944. The anatomy and embryology of the hemipenis of Lam- 
propeltis, Diadophis and Thamnophis and their value as criteria of re- 
lationship in the family Colubridae. Proc. Iowa Acad. Sci., vol. 51, 
pp. 411-445. 

Conant, R. 1963. Evidence for the specific status of the water snake Natrix 
fasciata. Amer. Mus. Novitates, no. 2122, pp. 1-38. 

Cope, E. D. 1900. The crocodilians, lizards, and snakes of North America. 
Ann. Rept. U. S. Nat. Mus., 1898, pp. 153-1294. 

Cowles, R. B., and C. M. Bogert. 1944. A preliminary study of the thermal 
requirements of desert reptiles. Bull. Amer. Mus. Nat. Hist., vol. 83, 
no. 5, pp. 265-296. 

Cowles, R. B., and G. L. Burleson. 1945. The sterilizing effect of high 
temperature on the male germ-plasm of the yucca night lizard, Xantusia 
vigilis. Amer. Nat., vol. 79, pp. 417-435. 

Davis, D. D. 1936. Courtship and mating behavior in snakes. Field Mus. 
Nat. Hist., Zool. Ser., vol. 20, no. 22, pp. 257-290. 

Dobzhansky, Th. 1951. Genetics and the origin of species. Columbia 
Univ. Press, New York, third ed. 

Dowling, H. G., and J. M. Savage. 1960. A guide to the snake hemipenis: 
a survey of basic structure and systematic characteristics. Zoologica, 
vol. 45, pt. 1, pp. 17-27. 

Edgren, R. A. 1952. A synopsis of the snakes of the genus Heterodon, with 
the diagnosis of a new race of Heterodon nasicus Baird and Girard. Nat. 
Hist. Misc., no. 112, pp. 1-4. 

Epling, C. 1947. Actual and potential gene flow in natural populations. 
Amer. Nat., vol. 81, pp. 104-113. 

Finneran, L. C. 1949. A sexual aggregation of the garter snake Thamnophis 
butleri (Cope). Copeia, 1949, no. 2, pp. 141-144. 

Fitch, H. S. 1956. Temperature responses in free-living amphibians and 
reptiles of northeastern Kansas. Univ. Kansas Publ., Mus. Nat. Hist., 
vol. 8, no. 7, pp. 417-476. 

Fox, W. 1954. Genetic and environmental variation in the timing of the 
reproductive cycles of male garter snakes. Jour. Morph., vol. 95, no. 3, 
pp. 415-450. 

Fox, W., and H. C. Dessauer. 1962. The single right oviduct and other 
urogenital structures of female Typhlops and Leptotyphlops. Copeia, 
1962, no. 3, pp. 590-597. 



346 Quarterly Journal of the Florida Academy of Sciences 

Haines, T. P. 1940. Delayed fertilization in Leptodeira annulata polysticta. 
Copeia, 1940, no. 2, pp. 116-118. 

Inger, R. F., and H. Marx. 1962. Variation of hemipenis and cloaca in the 
colubrid snake Calamaria lumbricoidea. Systematic Zool., vol. 11, no. 
1, pp. 32-38. 

Klauber, L. M. 1956. Rattlesnakes: their habits, life histories, and influ- 
ence on mankind. Univ. Calif. Press, Berkeley, 2 vols. 

Klinge, W. 1925. Kreuzung Zwischen Tropidonotus tesselatus = Mannchen 
X Tropidonotus viperinus = Weibchen. Bl. Aquar. Terrkde., vol. 36, 
pp. 20-21. 

Lederer, G. 1950. Ein Bastard von Elaphe guttata (Linne) = Mannchen 
X Elaphe quadrivittata quadrivittata (Holbrook) = Weibchen und 
dessen Riickkreuzung mit der miitterlichen Ausgangsart. Zool. Garten, 
vol. 17, pp. 235-242. 

Mayr, E. 1942. Systematica and the origin of species from the viewpoint of 
a zoologist. Columbia Univ. Press, New York. 

Mecham, J. S. 1961. Isolating mechanisms in anuran amphibians. In 
W. F. Blair (ed.), Vertebrate speciation, Univ. Texas Press, Austin, pp. 
24-66. 

Medsger, O. P. 1927. The hog-nosed snake or puff adder. Copeia, no. 
160, pp. 180-181. 

Mertens, R. 1950. C'ber Reptilienbastarde. Senckenbergiana, vol. 31, pp. 
127-144. 

. 1952. Kriechtiere und Lurche. Kosmos, Stuttgart. 



. 1956. tiber Reptilienbastarde, II. Senckenbergiana, vol. 37, pp. 

383-394. 

Mole, R. R. 1924. The Trinidad snakes. Proc. Zool. Soc. London, 1924, 
pp. 235-278. 

Munro, D. F. 1948. Mating behavior and seasonal cloacal discharge of a 
female Thamnophis sirtalis parietalis. Herpetologica, vol. 4, pp. 185- 
187. 

Neill, W. T. 1949. A new subspecies of rat snake (genus Elaphe), and 
notes on related forms. Herpetologica, vol. 5, 2nd suppl., pp. 1-12. 

. 1951. Notes on the natural history of certain North American 

snakes. Publ. Research Div., Ross Allen's Reptile Inst., vol. 1, no. 5, 
pp. 47-60. 

. 1962. The reproductive cycle of snakes in a tropical region, British 

Honduras. Quart. Jour. Florida Acad. Sci., vol. 25, no. 3, pp. 234-253. 



Neill: Isolating Mechanisms in Snakes 347 

. 1963. Polychromatism in snakes. Quart. Jour. Florida Acad. Sci., 

vol. 26, no. 2, pp. 194-216. 

. 1964. Taxonomy, natural history, and zoogeography of the rainbow 

snake, Farancia erytrogramma (Palisot de Beauvois). Amer. Midland 
Nat., vol. 71, no. 2, pp. 257-295. 

Noble, G. K. 1937. The sense organs involved in the courtship of Storeria, 
Thamnophis and other snakes. Bull. Amer. Mus. Nat. Hist., vol. 73, 
art. 7, pp. 673-725. 

Noble, G. K., and H. T. Bradley. 1933. The mating behavior of lizards; 
its bearing on the theory of sexual selection. Annals N. Y. Acad. Sci., 
vol. 35, pp. 25-100. 

Noble, G. K., and H. J. Clausen. 1936. The aggregation behavior of 
Storeria dekayi and other snakes, with especial reference to the sense 
organs involved. Ecol. Mono., vol. 6, no. 2, pp. 269-316. 

Pope, C. H. 1941. Copulatory adjustment in snakes. Field Mus. Nat. Hist., 
Zool. Ser., vol. 24, no. 22, pp. 249-252. 

SchOttler, W. H. A. 1950. Copula zwischen Bothrops und Crotalus. 
Deutsche Aquar. Terrar. Zeitschr., vol. 3, p. 14. 

Schweizer, H. 1941. Die Bastardform von Vipera aspis aspis X Vipera 
ammodytes ammodytes. Wochenschr. Aquar. Terrar. Kunde, vol. 38, 
pp. 345-346. 

Shaw, C. E. 1951. Male combat in American colubrid snakes with remarks 
on combat in other colubrid and elapid snakes. Herpetologica, vol. 7, 
no. 4, pp. 149-168. 

Smith, H. M. 1946. Hybridization between two species of garter snakes. 
Univ. Kansas Pub., Mus. Nat. Hist., vol. 1, no. 4, pp. 97-100. 

Woodbury, A. M. 1951. A snake den in Tooele County, Utah. Introduc- 
tion — a ten-year study. Herpetologica, vol. 7, pt. 1, pp. 4-13. 

Wright, A. H., and A. A. Wright. 1957. Handbook of snakes. Comstock 
Pub. Associates, Ithaca, N. Y., 2 vols. 

Florida State Museum and Department of Biology, University 
of Florida, Gainesville, Florida. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965) 



LARGE QUAHOG CLAMS FROM BOCA CIEGA BAY 
Harold W. Sims, Jr. 

On December 23, 1964, retired Air Force Major J. F. Simon 
brought to the Marine Laboratory a very large (168 mm, 6V2 lb.) 
southern quahog, Mercenaria campechiensis (Gmelin), found just 
off the intracoastal waterway at Redington Beach, in upper Boca 
Ciega Bay, Pinellas County, Florida. On December 29 three addi- 
tional large clams (149.5, 155.5, and 160 mm) were found buried 
in the substrate about one inch below the surface, with the siphon 
reaching the surface. 

The clams were found on a sand bar across the middle of a 
small bayou just west of the waterway. At normal low tide the 
water depth is 2-3 ft, but during spring low tides the bar is re- 
ported above water. It is composed of a mixture of sand, mud, 
and silt and is soft to a depth of about 1 ft, where hard bottom 
occurs. It is sparingly covered with manatee grass (Stjringodium) 
and turtle grass (Thalassia). On December 29 the water tempera- 
ture was 20 C and the salinity 32 0/00. 

It appears that three of these clams constitute new size records 
lor the species. Joseph Rosewater, Associate Curator of Mollusks, 
U. S. National Museum, states that the largest specimen in the col- 
lection measures ca. 153 mm and is the basis of the maximum size 
record for the species (Abbott, 1954). Porter and Chestnut (1960) 
list maxima of 5.5 inches (140 mm) and 4 lb. for this species in 
North Carolina. 

Literature Cited 

Abbott, R. T. 1954. American seashells. D. Van Nostrand Co., New York, 
514 pp. 

Porter, H. J., and A. F. Chestnut. 1960. The offshore clam industry of 
North Carolina. Proc. Nat. Shellfish Assn., vol. 51, pp. 67-73. 

Florida Board of Conservation Marine Laboratory, St. Peters- 
burg, Florida. Contribution No. 84. 

Quart. Jour. Florida Acad. Sci. 27(4) 1964 (1965): 348. 



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The next annual meeting will be at Tallahassee, March 11-13, 
1965. The chairman of the committee on local arrangements is 
Dr. Ruth S. Breen, Department of Botany, Florida State University. 



FLORIDA ACADEMY of SCIENCES 
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OFFICERS FOR 1964 

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Secretary: John D. Ktlby 

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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY OF SCIENCES 



VOLUME 28 



Editor 
Pierce Brodkorb 



Published by the 

Florida Academy of Sciences 

Gainesville, Florida 

1965 



Publication Dates of Volume 28 



Number 1 
Number 2 
Number 3 
Number 4 



May 12, 1965 
June 25, 1965 
September 22, 1965 
February 22, 1966 



Erratum 

No. 2, p. 226, line 21, read Mills for Miller. 



New Taxa Proposed in Volume 28 

\Palaeocarpilius brodkorbi Ross (Decapoda: Xanthidae) 236 

fKathpalmeria georgiana Ross (Cirripedia: Balanidae) 61, 63 

\Balanus (Armatobalanus) calvertensis Ross (Cirripedia: Balanidae) 334 

Bellator ribeiroi Miller (Pisces: Triglidae) 259 

\Leptodactylus abavus Holman (Amphibia: Leptodactylidae) 70 

Eleutherodactylus coqui Thomas (Amphibia: Leptodactylidae) 376 

Eleutherodactylus schwartzi Thomas (Amphibia: Leptodactylidae) 386 

\Rana miocenica Holman (Amphibia: Ranidae) 74 

f Rana bucella Holman (Amphibia: Ranidae) 76 

\Leiocephalus apertosulcus Etheridge (Reptilia: Iguanidae) 91 

Sphaerodactylus parthenopion Thomas (Reptilia: Gekkonidae) 117 

fHovacrex Brodkorb (Aves: Rallidae) 197 

f Idiornithidae Brodkorb (Aves: Gruiformes) 197 

fNeanis Brodkorb (Aves: Scytalopidae) ._, 197 



f Fossil 



CONTENTS OF VOLUME 28 



Number 1 

Radio astronomical observations from a canyon Heins Bollhagen, 

Jorge May, Jorge Levy, Thomas D. Carr, and Alex G. Smith 1 

Hawthorne, Bone Valley, and Citronelle sediments of Florida 

E. C. Pirkle, W. H. Yoho, and A. T. Allen 7 

A new cirriped from the Eocene of Georgia Arnold Ross 59 

Early Miocene anurans from Florida /. Alan Holman 68 

Fossil lizards from the Dominican Republic Richard Etheridge 83 

Sodium metaphosphate and mutton tenderness 

D. L. Huffman, A. Z. Palmer, J. W. Carpenter, and R. L. Shirley 106 

Bird remains from a Kentucky Indian midden Glen E. Woolfenden 115 

A new gecko from the Virgin Islands Richard Thomas 117 

Nereid blisters in Florida scallops Harry W. Wells 123 

Molybdenum toxicity in rats and rabbits 

L. R. Arlington, C. B. Ammerman, and J. E. Moore 129 



Number 2 

Oxidation of organic sulfides by dinitrogen tetroxide 

Robert D. Whitaker and Carole L. Bennett 137 

The phyllosoma larvae of Parribacus Harold W. Sims, Jr. 142 

Ciguatera poisoning from barracuda Edward Larson and Luis R. Rivas 173 

Young common snook on the coast of Georgia 

Thomas L. Linton and William L. Richards 185 

Food of Neoseps, the Florida sand skink 

Charles W. Myers and Sam R. Telford, Jr. 190 

Herring gulls diving for starfish 

Lowell P. Thomas and Shirley B. Thomas 195 

New taxa of fossil birds Pierce Brodkorb 197 

Ecology of the indigo bunting in Florida David W. Johnston 199 

Present status of the beaver in Florida 

James N. Layne and Bette S. Johns 212 

Dietary copper and enzymes in rabbit semen D. W. Stanley, 

R. L. Shirley, L. R. Arrington, and A. C. Warnick 221 

Florida Academy of Sciences award for 1965 225 

Florida Academy of Sciences charter and by-laws 226 



Number 3 

Notes of the Eocene Brachyura of Florida 

Jackson E. Lewis and Arnold Ross 233 

Confusion among species of Gomphus Minter J. Westfall, Jr. 245 

Florida fresh water fishes and conservation Luis R. Rivas 255 

A new species of searobin (Triglidae) George C. Miller 259 

Threadfin shad in Tampa Bay, Florida John H. Finucane 267 

Fossil birds from the Dominican Republic Lowell Bernstein 271 

Succinoxidase activity of the rat heart after whole-body gamma 

irradiation /. F. Easley, R. L. Shirley, and G. K. Davis 285 

Runway performance in two strains of rats 

Marshall B. Jones and Robert S. Fennell, III 289 

Experimental peritonitis and liver damage 

Steven I. Hajdu, Eva O. Hajdu, W. L. Holmes, and Alvan G. Foraker 297 

Ability of streams to assimilate wastes 

James B. Lackey and Hugh D. Putnam 305 

Officers and members of the Academy 318 



Number 4 

Oxidation of thianthrene by dinitrogen tetroxide 

Robert D. Whitaker and Carole L. Bennett 329 

Armatobalanus in the Miocene of Maryland Arnold Ross 332 

A Miocene needlefish from Bowden, Jamaica David K. Caldwell 339 

A huge Pleistocene box turtle from Texas /. Alan Holman 345 

Pleistocene lizards from New Providence Richard Etheridge 349 

Nuclear size in developing Acetabularia 

Kirsten R. Albrecht, Marinus J. Dijkman, and Manley L. Boss 359 

Feeding in the annelid Eteone heteropoda Joseph L. Simon 370 

Partial albinism in a blue crab 

Harold W. Sims, Jr., and Edwin A. Joyce, Jr. 373 

New species of Antillean Eleutherodactylus Richard Thomas 375 

Vitamin A and carotene in tissues of sheep R. L. Shirley, 

J. F. Easley, E. Sosa, T. J. Cunha, and A. C. Warnick 392 



•o in 3 

f- G ' ^ Quarterly Journal 

of the 

Florida Academy of Sciences 

Vol. 28 March, 1965 No. 1 



CONTENTS 

Radio astronomical observations from a canyon Heins Bollhagen, 

Jorge May, Jorge Levy, Thomas D. Can, and Alex G. Smith 1 

Hawthorne, Bone Valley, and Citronelle sediments of Florida 

E. C. Pirkle, W. H. Yoho, and A. T. Allen 7 

A new cirriped from the Eocene of Georgia Arnold Ross 59 

Early Miocene anurans from Florida /. Alan Holman 68 

Fossil lizards from the Dominican Republica Richard Etheridge 83 

Sodium metaphosphate and mutton tenderness 

D. L. Huffman, A. Z. Palmer, J. W. Carpenter, and R. L. Shirley 106 

Bird remains from a Kentucky Indian midden Glen E. Woolfenden 115 

A new gecko from the Virgin Islands Richard Thomas 117 

Nereid blisters in Florida scallops Harry W. Wells 123 

Molybdenum toxicity in rats and rabbits 

L. R. Arrington, C. B. Ammerman, and J. E. Moore 129 



Mailed May 12, 1965 

MM 1 9 



Quarterly Journal of the Florida Academy of Sciences 
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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY OF SCIENCES 



Vol. 28 March, 1965 No. 1 



RADIO ASTRONOMICAL OBSERVATIONS FROM A CANYON 

Heins Bollhagen, Jorge May, Jorge Levy, Thomas D. Carr, 
and Alex. G. Smith 

The portion of the electromagnetic spectrum which is access- 
ible to earth-bound radio astronomers is limited at its low-fre- 
quency end by the effects of the terrestrial ionosphere. Under 
good night-time observing conditions, reliable observations of ex- 
tra-terrestrial radio sources near the zenith can be made on a 
fairly large percentage of the nights at frequencies as low as 10 
Mc/s, and considerably less often at 5 Mc/s. However, even at 
times when ionospheric refraction and absorption of cosmic radio 
signals at these frequencies are not excessive, static from distant 
thunderstorms and interference from radio stations will often ob- 
scure the radiation under study. The interfering waves may be 
propagated around the surface of the earth from sources thousands 
of miles away by making one or more reflections from the iono- 
sphere, which is capable of reflecting such rays despite the fact 
that it is transparent when incidence is more nearly perpendicular. 
This work describes the results of a test which was made to de- 
termine the effectiveness of the walls of a deep canyon in shield- 
ing a low frequency radio telescope from low-angle terrestrial in- 
terference. 

Maipu Radioastronomical Orservatory 

The sources which have been studied most extensively at the 
lower frequencies by radio astronomers are the galactic source, the 
sun, and Jupiter (Smith and Carr, 1964). A substantial fraction of 
the data which are currently available on the low frequency noise 
bursts from Jupiter were obtained at the Maipu Radioastronomical 
Observatory, near Santiago, Chile (Carr et al., 1964). This ob- 
servatory is operated jointly by the University of Florida and the 



SMITHSONIAN 



SS^MAY181965 



2 Quarterly Journal of the Florida Academy of Sciences 

University of Chile. At the lowest frequencies at which the Jupiter 
observations have been made, 5 and 10 Mc/s, terrestrial interfer- 
ence has proven to be a serious problem. Good observing condi- 
tions are much rarer in this part of the spectrum than at the higher 
frequencies. 

The low-frequency antennas at the Maipu observatory are of 
relatively broad beam width, about 50° between half power points. 
Most of the rays comprising terrestrial interference arrive at the 
antenna at elevation angles of about 30° or less (see Ionospheric 
Radio Propagation, NBS Circular 462, U. S. Government Printing 
Office, 1949). Observations at the lower frequencies are most 
effective when the source is within about 45° of the zenith; how- 
ever, the antenna beams are so broad that even when they are di- 
rected vertically the interference entering from the sides is often 
excessive. 

It should be possible to improve observing conditions consider- 
ably by increasing the gain of the antenna system for radiation 
arriving from angles above 45° relative to its gain for low-angle 
radiation. The obvious solution, to build a more directive an- 
tenna, is not feasible because of the size of the structure which 
would be required. Not only would the main lobe of the antenna 
need to be sufficiently narrow, but effective supression of the side 
lobes would also be necessary. Regardless of the type of antenna 
used, whether it be a paraboloid or an array of smaller antennas, 
it would need to be of several wavelengths extent longitudinally 
and transversely (i.e., hundreds of feet). Tracking of the drifting 
cosmic sources would necessitate either the rotation of the struc- 
ture as a whole, or the continual readjustment of the phases of its 
elements. 

A more practicable solution would seem to be the use of a 
fixed broad-beam antenna with a barrier to shield it from the low- 
angle interference. To be effective, the barrier would need to be 
many wavelengths distant, and it should subtend an angle of 30 
to 45 degrees above the horizontal from the antenna. Shielding 
by such an obstacle would of course not be complete because of 
diffraction of radiation into the shadow zone; nevertheless, a con- 
siderable improvement in the ratio of the intensity of the desired 
signal to that of the interference would be expected. 

The Maipu observatory is located on a plain surrounded by 
hills, which unfortunately are too low to offer protection from 



Bollhagen et al.: Radioastronomical Observations 3 

terrestrial interference. However, innumerable valleys and can- 
yons abound in the nearby Andean foothills. It was therefore 
decided to install a radio telescope on the floor of a suitable can- 
yon, and to compare the terrestrial interference reaching this radio 
telescope with that experienced at the same time by an identical 
instrument on the exposed plain at Maipu. 

Peineta Canyon Installation 

With the aid of Dr. Jurgen Stock, director of the astronomical 
observatory being constructed at Cerro Tololo in the Chilean 
Andes under the auspices of Associated Universities for Research 
in Astronomy, a suitable canyon for the experiment was found near 
Cerro La Peineta, about 650 kilometers north of Maipu. A 10 
Mc/s antenna array, a battery-operated receiver, and a recorder 
were temporarily installed on the floor of the canyon. The an- 
tenna array, which was identical with the one at the same fre- 
quency at Maipu, consisted of two horizontal equi-phase full-wave 
dipoles a quarter wavelength above ground and a half wavelength 
apart. It was expected that most of the static would arrive from 
the equatorial jungle areas to the north and northeast; accordingly, 
the antenna was so situated that the shielding was greatest from 




Fig. 1. Profile of north wall of Peineta Canyon; antenna position in- 
dicated by A. 



4 Quarterly Journal of the Florida Academy of Sciences 

these directions. The profile of the north side of the canyon is 
shown in Figure 1. The top of this wall was about 50° above 
the horizontal when viewed from the antenna. The antenna lies 
well within the geometrical shadow of the north wall for most of 
the rays from distant sources of interference; nevertheless, some 
radiation will be diffracted around the barrier and will reach the 
antenna. The relative intensity of the diffracted radiation can 
readily be estimated with the aid of the Cornu Spiral (Jenkins and 
White, 1957). It was found in this way that for radiation arriving 
from the north at a 30° elevation angle, the intensity of the 
diffracted wave at the antenna was 13 decibels less than the in- 
tensity would have been if there had been no obstacle at all. 
Radiation from near the zenith, on the other hand, would be 
affected hardly at all by the walls of the canyon. Gross over- 



MAtPU tO Mc/s 

Dec. 17, 1962 



i u. 
! o 


( 

2345 


f 

GALACTIC NOiSE 

/- i- 1 , 

233C 


o 


f 

2315 


2300 


1 ~* 




■ •-.--• • \ rvw 








O 
a , 

1 O i_i 

a \ . 
a 


A 7^ 


PEINE t A canyon 
\ Dec. IT, ! 9 62 \ \J 












^M^u A udMW Mmk i 



GALACTIC JMO*St 



2345 2330 2315 230C 

*—— CHILiAN TIME 

Fig. 2. Records obtained simultaneously at the two stations. The spikes 
are static bursts. 



Bollhagen et al.: Radio astronomical Observations 5 

simplifications were of course made in arriving at the value of 
13 decibels; the north wall was assumed to be horizontal and per- 
pendicular to the ray path, and reflections from the other walls 
were neglected. Nevertheless, this preliminary calculation is in- 
dicative that a large gain in the ratio of the intensity of the desired 
signal to that of the interference can be achieved by using a nat- 
ural barrier for shielding. 

Results of the Experiment 

The simultaneous observations at Peineta Canyon and at Maipu 
were performed during December, 1962. The galactic noise com- 
ing from overhead was monitored, and the bursts of terrestrial in- 
terference appeared superimposed on the galactic background. 
Neither Jupiter nor the sun were in the antenna beams during the 
periods of measurement, and so far as is known, there are no other 
cosmic sources which radiate strongly enough to be detected with 
the equipment used. 

A typical pair of the simultaneously-obtained records is shown 
in Figure 2. The deflections resulting from the background of 
galactic noise were maintained at the same level at the two sta- 
tions by manual adjustment of the receiver gains. 

The period of simultaneous observation each night was from 
2000 to 0230 Chilean Time, except that on one night the period 
extended from 2000 to 0700. The total observing time was 61 
hours 35 minutes. 

Quantitative results are shown in Table 1. The quality of 
observing conditions was rated on a scale from zero to 5. A 
rating of 5 indicated that there was no interference whatsoever, 

TABLE 1 
Observing conditions at the two stations 



Quality of 














Duration 




observing 


Peineta Canyon 




Maipu 


conditions 












Hours 




Hours 


5 (perfect) 












15.0 




1.0 


4 












18.7 




1.9 


3 












11.8 




16.3 


2 or less 












16.1 




42.4 


Percentage 


of total observing 


time 








with good (4 


or 


5) 


conditions 




55% 




4.7% 



6 Quarterly Journal of the Florida Academy of Sciences 

while zero indicated that interference was at its worst. During 
zero conditions, for example, it would have been impossible to 
detect even the strongest noise bursts from Jupiter, had they been 
present. This rating scale is regularly used during the Jupiter 
watches at the University of Florida Radio Observatory and at the 
Maipu Radioastronomical Observatory. When conditions are 4 
or 5, even very weak Jupiter activity could be detected, while con- 
ditions of 2 or lower are generally considered to be unsatisfactory. 

Conclusions 

The results clearly indicate that for relatively low frequency 
observations of cosmic sources not far from the zenith, a consid- 
erable reduction in terrestrial interference can be achieved by 
placing the radio telescope on the floor of a deep canyon. The 
use of naturally shielded sites should make possible the extension 
of Jupiter's decametric spectrum to frequencies somewhat lower 
than the present limit, and should result in improved observing 
conditions at the higher frequencies as well. 

Acknowledgments 

We wish to thank Dr. Jurgen Stock of the Associated Universi- 
ties for Research in Astronomy for his assistance in finding the site 
for this experiment and for providing many of the necessary facili- 
ties. We also thank Sr. Juan Aparici for his helpful assistance in 
making the observations, and the Observatorio Astronomico Nac- 
ional of Chile for providing facilities and support. We gratefully 
acknowledge the financial suport of the U. S. National Science 
Foundation, the Office of Naval Research, and the Army Research 
Office (Durham). 

Literature Cited 

Carr, T. D., G. W. Brown, A. G. Smith, C. S. Higgins, H. Bollhagen, 
J. May, and J. Levy. 1964. Spectral distribution of the decametric 
radiation from Jupiter in 1961. Astrophysical Journal, vol. 140, no. 2, 

pp. 778-795. 

Jenkins, Francis A., and Harvey E. White. 1957. Fundamentals of op- 
tics. McGraw-Hill Book Co., New York, 637 pp. 

Smith, Alex. G., and Thomas D. Carr. 1964. Radio exploration of the 
planetary system. D. Van Nostrand Co., Princeton, N. J., 148 pp. 

Quart. Jour. Florida Acad. Sci. 28(1) 1965 



HAWTHORNE, BONE VALLEY, AND CITRONELLE 
SEDIMENTS OF FLORIDA 

E. C. Pirkle, W. H. Yoho, and A. T. Allen 

During the course of several years of geological work in Flor- 
ida many sections of sediments have been measured, samples col- 
lected, and the materials analyzed in detail. Such data are of in- 
estimable value in formulating geological ideas, in testing geologi- 
ical concepts, and in executing exploration work for economic 
mineral occurrences. The purpose of this report is to make avail- 
able some of the detailed sections and analyses and to indicate 
possible applications of the data to current geological problems. 
The sections selected for presentation are of Hawthorne, Bone 
Valley, and Citronelle materials. 

Hawthorne Formation 

The Hawthorne Formation is present either at the surface or 
in the subsurface throughout most of peninsular Florida. The 
formation is absent only in the area of the Ocala uplift in the 
west-central part of the peninsula. Even there, however, erosional 
remnants of the Hawthorne are present. 

The main body of the Hawthorne Formation is considered by 
recent workers to be of early Miocene age or of early and middle 
Miocene age (Brodkorb, 1963, p. 159; Bishop, 1956, p. 25; and 
Espenshade and Spencer, 1963, p. 26). It consists of clays, sands, 
dolomites, and limestones (both calcic and dolomitic). Phosphate 
particles are present in the sediments. Locally this phosphorite 
is one of the dominant constituents of the beds. In some localities 
extensive phosphate concentrations occur at the top of the main 
body of the Hawthorne Formation. These accumulations consist 
of pebbles and grains of phosphorite embedded in a matrix that 
may be nearly all quartz sand or clay or carbonate but that usually 
is a mixture of these sediments. The concentrations differ in 
origin and in age. 

Descriptions of Hawthorne Sediments 

A number of problems are encountered in naming and de- 
scribing some of the units of the Hawthorne Formation. Units 



8 Quarterly Journal of the Florida Academy of Sciences 

composed almost entirely of clay or sand or carbonate cause no 
particular difficulty. However, many Hawthorne units consist of 
varying combinations of quartz sand, silt, clay, finely divided car- 
bonate (both calcic and dolomitic), detrital pebbles and blocks 
of limestones or marls of varying purity, sand-size to pebble-size 
phosphorite, detrital pebbles and blocks of clay, and other con- 
stituents. In addition, the various combinations of these sediments 
usually change considerably in short distances, both vertically 
and laterally. 

Estimating quantities of these mixtures of sediments from field 
observations is not satisfactory. Experience has shown that such 
field estimates can be quite unrealistic. Nor will the naming of 
some of these combinations on the sole basis of the break-down 
of the sediments into such fractions as the percentages of clay, 
silt, sand, granules, pebbles, and soluble materials always give a 
true impression as to the nature of the units. 

In this work the major concern in describing the Hawthorne 
units has been to accurately portray the nature of the sediments. 
In cases this can be accomplished by using names such as clay and 
limestone. In other cases where a specific name might be mis- 
leading, only a description believed representative of the unit is 
given. 

Analyses of Hawthorne Sediments 

In analyzing materials collected from the Hawthorne Forma- 
tion, the sediments selected were continually and carefully halved 
until the desired representative fraction needed for analysis was 
obtained. This representative sample was then divided into four 
parts. One part was treated with hydrochloric acid to obtain 
the per cent insoluble residue and the per cent soluble. The in- 
soluble residue portion was carefully screened and each mesh 
size examined under a binocular microscope. The heavy minerals 
were separated by means of tetrabrom ethane from the 1/16 to 1/8 
mm size material, mounted into slides, and in most cases between 
300 and 400 grains were identified and counted. The relative 
abundance of the heavy minerals is given in various tables. The 
portion of the insoluble residue that passed through a 325 mesh 
screen was considered as clay. The results of the insoluble resi- 
due analyses are given in table form. 

A second portion of the original quartered sample was water- 



Pirkle, Yoho, Allen: Analyses of Sediments 9 

washed and broken down into various mesh sizes. Each mesh 
size was analyzed under a binocular microscope. The results of 
these analyses are not presented in table form in this report but are 
utilized in combinations with the insoluble residue analyses and 
field observations in naming and describing the various units 
given in the sections. In addition, an occasional sample of the 
1/16 to 1/8 mm fraction was treated with acid to remove phos- 
phate and other solubles. The heavies were then extracted, identi- 
fied, and counted as a check against the heavy mineral counts ob- 
tained from the same size fraction of the insoluble residues. 

A third portion of the original quartered sample was utilized 
for the determination of the P 2 5 content of the sample. The 
P 2 5 analyses were run by Thornton and Company of Tampa, 
Florida, and are listed in the various tables of the report. 

The fourth part of the original quartered sample was treated 
with acid to remove phosphate and other soluble materials. The 
heavy minerals of this residue were extracted to determine the 
total per cent heavies of the materials. This value is listed in the 
tables. 

The above procedures in analyzing the samples were carried 
cut for all units presented in the Hawthorne sections, i.e. the 
Devil's Mill Hopper and Brooks Sink sections. The same meth- 
ods of treatment were followed in analyzing sediments from the 
Bone Valley district, i.e. the Peace River and Achan sections. In 
part the procedures for analyzing the sediments were selected 
and followed to provide specific data needed for future studies. 

Procedure in Treatment of Heavy Minerals 

Although individual counts were made of all heavy minerals 
selected for examinations and percentages were computed, the 
results are not given in percentage form; instead a system is used 
to present the data so that only relatively large differences are 
obvious (see Table 3). This procedure is followed because it is 
believed that field and laboratory techniques, although carried 
out with care, are not sufficiently accurate to justify conclusions 
based on small differences, either percentage-wise or mineral- 
wise. 

It is well known that heavy mineral frequencies are different 
for various size fractions of any given sample of sediments. The 



10 Quarterly Journal of the Florida Academy of Sciences 

TABLE 1 

Devil's Mill Hopper, Alachua County, Florida (Pirkle et al., 1964) 
Surface sands and Hawthorne sediments * 







Insoluble 


residue 


Total 








Quartz 


Clay 


Total 






sand ( 


—325 mesh) 


soluble 


p 2 o 5 


heavies 


Sample 


Unit 


in % 


in % 


in % 


in % 


in % 


1 


16 


93.28 


3.70 


3.00 


0.92 


.14 


2 


15 


33.89 


8.77 


54.22 


15.49 


.17 


3 


15 


33.61 


22.71 


41.21 


12.76 


.15 


4 


15 


57.22 


21.43 


20.53 


5.78 


.30 


5 


14 


4.84 


2.02 


93.28 


1.45 


.04 


6 


13 


52.61 


44,39 


1,37 


0.85 


.09 


7 


13 


74.55 


21.00 


3.88 


1.05 


.22 


8 


13 


12.48 


12.40 


75.12 


0.51 


.03 


9 


12 


64.19 


28.13 


6.06 


3.14 


.24 


10 


12 


57.92 


32.55 


7.14 


2.05 


.13 


11 


12 


71.68 


22.46 


4.93 


2.42 


.18 


12 


12 


60.43 


28.04 


10.90 


4.41 


.16 


13 


11 


38.73 


29.15 


31.95 


8.86 


.12 


14 


11 


51.88 


27.20 


20.97 


8.90 


.15 


15 


10 


55.63 


29.64 


12.70 


7.51 


.21 


16 


10 


35.93 


29.73 


34,33 


7.99 


.25 


17 


10 


30.48 


50.51 


19.04 


3.08 


.15 


18 


9 


1.73 


69.27 


29.02 


1.46 


.01 


19 


8 


68.28 


20.46 


11.11 


3.53 


.20 


20 


7 


20.19 


23.26 


56,34 


1.89 


.11 


21 


6 


1.21 


39.10 


59.52 


0.99 


.02 


22 


6 


trace 


86.10 


14.00 


1.03 


.01 


23 


5 


23.78 


23.76 


52.41 


3.64 


.08 


24 


5 


25.04 


34.81 


38.04 


5.82 


.04 


25 


5 


38.60 


26.52 


34.69 


2.50 


.24 


26 


5 


55.11 


9.28 


35.41 


2.37 


.29 


27 


5 


46.23 


13.97 


39.40 


2.26 


.27 


28 


4 


36.86 


37.43 


25,32 


4.03 


.15 


29 


4 


26.85 


48.75 


23.85 


4.05 


.16 


30 


3 


1.22 


29.29 


68.97 


1.38 


.002 


31 


2 


55.50 


4.18 


40.28 


3.45 


.25 


32 


1 


21.26 


4.99 


73,50 


1.01 


.14 



* The column headings in this table are partly self-explanatory. How- 
ever, a few remarks are necessary. The value listed under the insoluble resi- 
due as the per cent quartz sand is the percentage of the original sample con- 
sisting of quartz sand. The per cent clay represents that part of the original 
sample consisting of insoluble sediments that passed through a 325 mesh 
screen. A part of this value includes silt-size materials; however, most of the 
—325 mesh sediments are of clay size. The percentage figure given under the 
heading of "total soluble" reflects mainly carbonate and phosphate content of 



Pirkle, Yoho, Allen: Analyses of Sediments 11 

particular heavy mineral fraction or fractions from which heavy 
minerals are separated and counted depend upon the objectives 
of the work. For this report, the heavy minerals counted were 
those separated from the 1/16 to 1/8 mm size-grade materials. 
Heavies from this fraction have value for correlation purposes. 
Although not utilized directly in the study, the total heavies for 
each sample were also examined. 

Because of the particular size-grade of sediments selected from 
which to analyze heavy minerals, the results of this work can not 
be directly compared to other published data on the heavy min- 
erals of Florida. For example, the counts given by Thoenen and 
Warne (1949) were for total heavies; the data given by Carr and 
Alverson (1959) were for all heavies that would not pass through 
a 100 mesh screen. In the present work some of the heavies that 
occur in only minute amounts were not counted. Such disre- 
garded heavy minerals include monazite, spinel, corundum, sphene, 
and zoisite. 

Even though the utmost care in sampling and in making lab- 
oratory analyses is practiced, human errors may occur in studying 
such sediments as those comprising the Hawthorne, Bone Valley, 
and Citronelle materials. In order to reduce such variations and 
to make for more uniformily prepared data, the same person car- 
ried out the same assignment throughout the entire study. Pirkle 
was responsible for the field work, Yoho for all laboratory anal- 
yses, and Allen for the identification and counting of all heavy 
minerals. 

Devil's Mill Hopper (Cotype Locality) 

A number of sections of the Miocene Hawthorne Formation 
have been measured by different workers at the Devil's Mill Hop- 
per, a large sink hole located about 6 miles northwest of Gaines- 
ville in Alachua County. These sections, based largely on visual 
field observations, include those reported by Cooke (1945, p. 147); 
Pirkle (1956, p. 215; 1958, p. 150); Puri, Bishop, and Stewart (Puri 
and Vernon, 1959, p. 122); and Thompson and Floyd (McClellan, 
1962). The sections differ in various aspects, a result to be ex- 

the sediments. The P2O5 value is the percentage of P 2 5 in the complete 
sample. The total per cent heavies is the percent of the sample or unit con- 
sisting of heavy minerals. This value, however, does not include phosphorite, 
mica, or iron concretions. Sample and unit numbers correspond to those of 
the measured section (see Appendix). 



12 Quarterly Journal of the Florida Academy of Sciences 



TABLE 2 

Devil's Mill Hopper, Alachua County, Florida 
Mechanical analyses of quartz sand fraction of various units 







Quartz 
















sand 


Per cent total quartz 


sand retained on 


mesh 


Sample 


Unit 


in % 


10 18 


35 


60 


120 


230 


1 


16 


93.28 


.04 


4.30 


42.40 


40.51 


8.46 


2 


15 


33.89 




1.73 


15.95 


77.10 


4.83 


3 


15 


33.61 




1.06 


24.11 


70.47 


3.92 


4 


15 


57.22 




2.16 


26.49 


51.83 


18.84 


5 


14 


4.84 




7.49 


52.42 


36.56 


3.08 


6 


13 


52.61 


,30 


10.00 


56.74 


23.11 


8.00 


7 


13 


74.55 


.23 


7.70 


44.59 


35.83 


11.19 


8 


13 


12.48 


.16 


10.08 


56.16 


21.60 


10.08 


9 


12 


64.19 


.03 .29 


9.10 


56.65 


23.91 


8.59 


10 


12 


57.92 


.29 


10.30 


57.19 


22.55 


8.03 


11 


12 


71.68 


,30 


9.06 


58.01 


24.42 


8.21 


12 


12 


60.43 


.39 


8.97 


58.34 


24.82 


7.48 


13 


11 


38.73 


.46 


10.24 


66.46 


18.66 


3.71 


14 


11 


51.88 


.29 


11.76 


62.50 


19.37 


5.53 


15 


10 


55.63 


.75 


10.80 


54.56 


25.49 


7.73 


16 


10 


35.93 


trace 


2.82 


29.88 


24.18 


43.12 


17 


10 


30.48 




25.12 


56.57 


13,31 


4.18 


18 


9 


1.73 


trace 


3.83 


4.88 


4.53 


86.76 


19 


8 


68.28 


trace 


3,37 


49,59 


46.13 


.91 


20 


7 


20.19 




3.41 


35.13 


59.10 


1.35 


21 


6 


1.21 




trace 


40.91 


42.42 


16.67 


22 


6 


trace 












23 


5 


23.78 




5.50 


57.92 


31.58 


4.83 


24 


5 


25.04 






.15 


1.24 


97.54 


25 


5 


38.60 






8.22 


86.54 


4.73 


26 


5 


55.11 




.04 


8.90 


81.12 


9.58 


27 


5 


46.23 






8.87 


82.64 


8.06 


28 


4 


36.85 




.16 


26.05 


69.42 


3.29 


29 


4 


26.86 




1.01 


29.70 


63.22 


4.12 


30 


3 


1.22 






11.48 


8.20 


80.33 


31 


2 


55.50 




1.15 


23.87 


69.91 


5.00 


32 


1 


21.26 




.66 


33.80 


52.81 


11.61 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 13 

pected because of the lens-like nature of the Hawthorne sedi- 
ments and the rapid changes, even within individual small lenses, 
in the proportions of carbonate, quartz sand, clay, and phosphate 
particles. Furthermore, numerous landslides and slow slumpage 
at the sink result in the continual exposure of new lenses of differ- 
ent type sediments at the same elevation where other lenses had 
previously been recorded in sections. While collecting samples 
to make the heavy mineral counts given in this report, a new sec- 
tion showing the presently exposed strata was measured. As an 
aid in describing materials in this new section (see Appendix) 
numerous laboratory analyses were made and used to supplement 
field observations. Some of these analyses are given in Tables 
1 and 2. 

Lithologic characteristics of the sediments at the Mill Hopper 
sink have been discussed previously (Pirkle, 1958). These features 
include silicified clay; the gradation of clay into lenses and irreg- 
ular masses of limestone and sandstone; the apparent replacement 
of clay by carbonate; intraformational breccias and conglomerates; 
and a phosphate reaction zone showing limestone replacement re- 
sulting in "hard-rock" type phosphate. McClellan (1962) studied 
the clay minerals in the Devil's Mill Hopper by means of X-ray 
diffraction methods and the electron microscope. Isphording (1963) 
treated the heavy minerals in the sediments. The results of Is- 
phording's work and the results presented in this report (Table 3) 
were arrived at independently and are comparable. 

Of interest in regard to the heavy minerals is the persistence 
of garnet throughout the Hawthorne sediments. Garnet is present 
in the phosphate concentrations at the top of the main body of the 
Hawthorne Formation (samples 2-4, Table 3). Likewise the min- 
eral is present in limestones (samples 5, 8, and 32), in clays (sample 
22), in quartz sands, and in all of the various combinations of these 
sediments. Also of interest is the presence locally of considerable 
amounts of epidote. At the Mill Hopper the upper Hawthorne 
materials are relatively high in this mineral (samples 2-5, Table 3). 
As will be noted from tables given throughout this report, Haw- 
thorne and Bone Valley sediments are characterized by the per- 
sistence of garnet and the presence locally of relatively large 
amounts of epidote. In contrast, garnet is essentially absent in 
Citronelle sediments, and epidote, if present, occurs in relatively 
minute amounts. 



14 Quarterly Journal of the Florida Academy of Sciences 

TABLE 3 

Devil's Mill Hopper, Alachua County, Florida 
Heavy minerals in 1/16 to 1/8 mm fraction * 



-2* 

s 


*3 


•a 




a 

03 

X 

o 
o 

a> 
h4 


e 
o 

g 

S3 


W58 


03 

""3 
M 

C/3 


a 

Is 

o 
Eh 


03 
O 
13 
'ft 

w 


03 

s 

o 


1 


16 


vc 


c 


VC 


vc 


vc 


R 


R 


_ 


VR 


2 


15 


c 


c 


c 


c 


c 


R 


VR 


A 


R 


3 


15 


vc 


R 


c 


c 


c 


R 


R 


VC 


C 


4 


15 


A 


C 


c 


R 


c 


R 


R 


VC 


R 


5 


14 


A 


C 


R 


vc 


c 


R 


VR 


C 


C 


6 


13 


A 


C 


VC 


c 


R 


C 


R 


R 


VR 


7 


13 


VA 


R 


C 


c 


R 


C 


R 


R 


R 


8 


13 


VC 


C 


A 


c 


R 


R 


R 


R 


VR 


9 


12 


A 


R 


C 


vc 


R 


C 


C 


R 


R 


10 


12 


A 


R 


C 


vc 


R 


C 


C 


R 


VR 


11 


12 


A 


R 


C 


vc 


R 


C 


R 


R 


R 


12 


12 


A 


R 


C 


c 


R 


C 


R 


R 


R 


13 


11 


VA 


R 


C 


c 


C 


C 


R 


R 


R 


14 


11 


A 


R 


C 


vc 


C 


C 


R 


R 


R 


15 


10 


VA 


C 


C 


vc 


R 


C 


R 


R 


R 


16 


10 


VA 


R 


C 


c 


R 


C 


R 


R 


R 


17 


10 


VA 


R 


C 


vc 


R 


C 


R 


R 


C 


18 


9 


A 


C 


VC 


c 


C 


C 


VR 


R 


C 


19 


8 


VA 


C 


c 


vc 


R 


C 


VR 


R 


c 


20 


7 


A 


R 


c 


vc 


R 


C 


VR 


R 


c 


21 


6 


A 


C 


c 


vc 


VR 


C 


R 


R 


c 


22 


6 


VA 


R 


c 


vc 


R 


R 


VR 


VR 


c 


23 


5 


VA 


C 


c 


c 


R 


C 


R 


R 


c 


24 


5 


VA 


R 


vc 


c 


VR 


C 


R 


R 


c 


25 


5 


VA 


R 


c 


vc 


VR 


R 


R 


R 


c 


26 


5 


A 


C 


c 


vc 


VR 


C 


R 


R 


c 


27 


5 


A 


C 


c 


c 


R 


C 


R 


R 


c 


28 


4 


A 


C 


c 


vc 


R 


c 


R 


R 


c 


29 


4 


A 


C 


c 


vc 


VR 


c 


R 


R 


c 


30 


3 


A 


C 


c 


vc 


VR 


R 


R 


C 


c 


31 


2 


A 


C 


c 


c 


R 


C 


R 


R 


c 


32 


1 


A 


C 


c 


c 


R 


R 


R 


R 


c 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 

In listing the relative abundance of the different heavy minerals the 
following scale was used: 



Pirkle, Yoho, Allen: Analyses of Sediments 15 

Brooks Sink (Cotype Locality) 

The best exposures of the phosphatic Miocene Hawthorne For- 
mation are at Brooks Sink, a large sink hole located in southern 
Bradford County about 4 miles east of the town of Brooker. There 
approximately 40 feet of Hawthorne sediments are exposed and 
are overlain by shell marl. This shell marl has been dated by 
Puri on the basis of ostracods (Pirkle, 1956, p. 210) as lower Choc- 
tawhatchee in age (late middle Miocene or late Miocene). The 
exposures at this sink offer an unusually good opportunity to study 
various types of phosphate particles and the occurrences of various 
types of concentrations of phosphorite. The analyses of the size 
distribution of the quartz grains in phosphate particles, in the 
matrix containing the phosphate particles, and in surrounding beds 
are of interest in these studies. 

The same kinds of sedimentary features observed at Brooks 
Sink have been seen at some exposures where phosphatic sediments 
were being mined in the Bone Valley district. Sections and de- 
scriptions of the sink have been published by Sellards (1909, p. 
240), Pirkle (1956, p. 207), Puri and Bishop (Puri and Vernon, 1959, 
p. 120), and Espenshade and Spencer (1963, p. 75). Espenshade 
and Spencer give fossil data and phosphate analyses with their 
section. In all of the sections the combinations of sediments pres- 
ent in each unit or bed were described for the most part from 
field observations. The new section given in the Appendix of this 
report is based on both field observations and laboratory analyses. 
Tables 4-6 illustrate features of the various units exposed at 
this sink. 

Slight differences in heavy mineral occurrences can be noted 
from Table 6. For example, beginning with unit 6 (sample 25) 
and going downward through unit (sample 33) ilmenite and 



Greater than 75% Flood _F 

50 to 75% Very Abundant VA 

25 to 50% Abundant A 

15 to 25% Very Common VC 

5 to 15% Common C 

1 to 5% Rare R 

Less than 1% Very Rare VR 

To illustrate, if 20% of the heavy mineral particles extracted from the 
1/16 to 1/8 mm fraction of sediments consisted of ilmenite, the relative il- 
menite content was considered as very common and "VC" was entered for 
that particular heavy mineral. Data for only the more common heavy min- 
erals are given. 



16 Quarterly Journal of the Florida Academy of Sciences 

TABLE 4 

Brooks Sink, Bradford County, Florida 
Surface sands, Late Miocene and Hawthorne sediments * 







Insoluble 


residue 


Total 








Quartz 


Clay 


Total 






sand (- 


-325 mesh) 


soluble 


p 2 o 5 


heavies 


Sample 


Unit 


in % 


in % 


in % 


in % 


in % 


1 


20 


93.48 


4.97 


1.75 


0.25 


.28 


2 


20 


79.76 


18.48 


2.00 


0.13 


.14 


3 


19 


1.68 


1.10 


97.12 


0.15 


.02 


4 


19 


6.16 


2.61 


91.16 


0.55 


.10 


5 


19 


21,38 


10.52 


67.96 


1.60 


.28 


6 


18 


59.25 


3.70 


36.87 


1.41 


.60 


7 


18 


58.06 


5.91 


36.04 


1.48 


.61 


8 


18 


58.07 


5.64 


36.28 


1.64 


.84 


9 


17 


13.73 


8.25 


77.97 


1.82 


.12 


10 


16 


27.32 


6.48 


66.16 


6.41 


.23 


11 


15 


7.96 


3.10 


88.90 


1.36 


.04 


12 


14 


18.51 


3.54 


77.77 


22.16 


.15 


13 


14 


24.76 


5.83 


69.30 


3.87 


.15 


14 


13 


4.82 


3.80 


91.35 


3.37 


.03 


15 


12 


19.24 


4.46 


76,30 


5.58 


.16 


16 


12 


23.74 


15.66 


59.94 


18.10 


.15 


17 


11 


28.60 


8.14 


63.22 


2.83 


.06 


18 


10 


8.43 


4.62 


86.93 


1.65 


.41 


19 


9 


25.86 


9.37 


64.61 


2.88 


.14 


20 


9 


42.79 


8.52 


48.65 


4.16 


.25 


21 


9 


31.91 


8.98 


59.08 


3.70 


.17 


22 


8 


20.24 


5.22 


74.31 


1.75 


.08 


23 


7 


50.24 


8.57 


41.15 


4.65 


.12 


24 


7 


47.74 


10.77 


41.41 


4.84 


.14 


25 


6 


6.72 


6.38 


86.75 


2.41 


.06 


26 


5 


41.12 


12.12 


46.53 


5.35 


.29 


27 


4 


22.57 


10.68 


66.67 


13.94 


.36 


28 


4 


.96 


5.01 


93.99 


2.21 


.04 


29 


3 


26.00 


11.40 


62.40 


11.81 


.43 


30 


2 


1.10 


8.67 


89.45 


0.89 


.02 


31 


1 


6.63 


60.90 


30.66 


2.58 


.06 


32 


1 


23.30 


13.71 


62.49 


16.18 


.19 


33 





34.79 


6.25 


58.85 


2.77 


.20 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 



17 



TABLE 5 

Brooks Sink, Bradford County, Florida 
Mechanical analyses of quartz sand fraction of various units 







Quartz 
















sand 


Per cent total quartz 


sand retained on 


mesh 


Sample 


Unit 


in % 


10 18 


35 


60 


120 


230 


1 


20 


93.48 


1.93 


17.12 


35.16 


30.93 


14.86 


2 


20 


79.76 


0.04 6.03 


30.61 


47.88 


12.49 


2.94 


3 


19 


1.68 






8.33 


78.57 


13.10 


4 


19 


6.16 






1.63 


80.13 


18.24 


5 


19 


21.38 






2.52 


93.56 


3.92 


6 


18 


59.25 






1.82 


87.12 


11.06 


7 


18 


58.06 




.10 


1.41 


79.00 


19.48 


8 


18 


58.07 




.03 


1.85 


83.69 


14.42 


9 


17 


13.73 




.29 


3.93 


72.49 


22.85 


10 


16 


27.32 




1.46 


13.12 


75.73 


9.62 


11 


15 


7.96 






12.53 


78.45 


8.77 


12 


14 


18.51 




1.62 


21.17 


66.74 


10.48 


13 


14 


24.76 




.48 


11.65 


83.13 


4.74 


14 


13 


4.82 




.41 


20.75 


67.63 


11.20 


15 


12 


19.24 




.31 


12.79 


83.26 


3.64 


16 


12 


23.74 


.42 


6.06 


27.95 


59.43 


6.14 


17 


11 


28.60 




.28 


11.65 


82.78 


5.30 


18 


10 


8.43 




.95 


8.06 


83.41 


7.58 


19 


9 


25.86 




1.01 


13.66 


80.09 


5.23 


20 


9 


42.79 


.05 


.84 


17.54 


76.43 


5.15 


21 


9 


31.91 


.13 


.69 


20.61 


74.00 


4.57 


22 


8 


20.24 




1.09 


22.43 


64.33 


12.15 


23 


7 


50.24 




2.95 


37.66 


57.83 


1.55 


24 


7 


47.74 


.08 


3.32 


35.99 


58.84 


1.76 


25 


6 


6.72 




5.65 


43.45 


42.56 


8.33 


26 


5 


41.12 




4.57 


25.64 


62.51 


7.29 


27 


4 


22.57 


4.95 


24.76 


36.16 


31.65 


2.48 


28 


4 


.96 


6.25 2.08 


18.75 


33.33 


31.25 


8.33 


29 


3 


26.00 


1.38 2.69 


17.08 


39.31 


37.92 


1.62 


30 


2 


1.10 




3.64 


5.45 


7.27 


83.64 


31 


1 


6.63 






1.50 


3.00 


95.50 


32 


1 


23.30 


1.11 4.36 


11.04 


32.34 


45.77 


5.39 


33 





34.79 


.17 


3.15 


31.31 


63.25 


2.12 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



18 Quarterly Journal of the Florida Academy of Sciences 



TABLE 6 

Brooks Sink, Bradford County, Florida 
Heavy minerals in 1/16 to 1/8 mm fraction * 



2 

C/3 


4-> 

*2 


*2 




a 


o 
o 


a 
o 
o 
u 


a> '2 
II 


1 


o 

Eh 


o 

12 
a 

w 


C 

S-H 


1 


20 


A 


c 


VC 


VC 


C 


VR 


VR 








2 


20 


A 


c 


c 


VC 


C 


C 


VR 


— 


VR 


3 


19 


A 


c 


R 


R 


VC 


R 


R 


A 


R 


4 


19 


VC 


R 


R 


R 


c 


R 


R 


A 


R 


5 


19 


VC 


R 


VR 


R 


c 


R 


R 


A 


R 


6 


18 


VC 


R 


R 


R 


VC 


R 


R 


A 


R 


7 


18 


A 


R 


R 


C 


VC 


R 


R 


A 


R 


8 


18 


VC 


R 


R 


R 


c 


R 


R 


A 


R 


9 


17 


A 


R 


R 


C 


VC 


R 


R 


A 


R 


10 


16 


VC 


R 


R 


C 


VC 


R 


R 


A 


R 


11 


15 


A 


R 


R 


C 


c 


VR 


VR 


A 


R 


12 


14 


A 


R 


R 


C 


c 


VR 


VR 


A 


C 


13 


14 


A 


R 


R 


R 


c 


VR 


VR 


A 


R 


14 


13 


A 


R 


R 


C 


VC 


R 


VR 


VC 


R 


15 


12 


A 


R 


R 


C 


VC 


R 


VR 


VC 


R 


16 


12 


A 


R 


R 


c 


VC 


R 


R 


c 


R 


17 


11 


VC 


R 


R 


R 


A 


C 


R 


c 


R 


18 


10 


A 


R 


R 


C 


VC 


R 


R 


c 


C 


19 


9 


A 


R 


R 


R 


VC 


R 


R 


c 


R 


20 


9 


A 


VR 


R 


R 


VC 


C 


R 


c 


R 


21 


9 


A 


R 


R 


R 


VC 


C 


R 


c 


R 


22 


8 


A 


R 


R 


C 


VC 


C 


R 


c 


R 


23 


7 


A 


R 


R 


C 


c 


R 


R 


VC 


R 


24 


7 


A 


R 


R 


R 


VC 


C 


R 


VC 


R 


25 


6 


A 


R 


C 


VC 


c 


R 


VR 


R 


R 


26 


5 


A 


R 


R 


C 


c 


C 


R 


VR 


R 


27 


4 


A 


C 


R 


VC 


c 


C 


VR 


VR 


R 


28 


4 


A 


C 


R 


VC 


R 


R 


R 


R 


R 


29 


3 


VA 


R 


R 


C 


c 


C 


R 


— 


R 


30 


2 


A 


R 


R 


C 


c 


C 


R 


— 


R 


31 


1 


VA 


R 


R 


C 


R 


c 


R 


— 


R 


32 


1 


VA 


R 


R 


C 


c 


c 


R 


VR 


R 


33 





VA 


R 


R 


C 


c 


c 


VR 


— 


C 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 19 

zircon are relatively more abundant and kyanite-sillimanite and 
epidote are more scarce. However, of more significance are the 
occurrences of garnet in the Hawthorne sediments, and the pres- 
ence locally of relatively substantial amounts of epidote. As in 
the case of the Devil's Mill Hopper the garnet is present through- 
out all of the Hawthorne sediments regardless of their composi- 
tion; units containing a high content of carbonate are character- 
ized by garnet (samples 14, 18, 28, and 30) as are those units con- 
taining a relatively high content of quartz sand (samples 20, 23, 
and 24) or clay (sample 31). Also of interest are the similarities 
of the heavy mineral suites of the Hawthorne sediments and the 
late Miocene shell marl (unit 19), as illustrated by the occurrences 
of epidote and garnet. It has been noted in other areas, for 
example at some localities along the east side of the Lake Wales 
Ridge, that heavy mineral suites of late Miocene shell beds are 
similar to those of underlying Hawthorne materials. Likewise in 
some areas the contact relations between Hawthorne sediments 
and late Miocene shell beds are suspected to be gradational (Pirkle, 
1956, p. 210; Klein et al., 1964, p. 23). 

Phosphatic Sediments of the Bone Valley District 

The phosphatic sediments of the Bone Valley district of Polk, 
Hillsborough, Hardee, and Manatee counties are generally dated 
either as Miocene or as Pliocene in age (Simpson, 1929; Vernon, 
1951; Brodkorb, 1955; Cathcart, 1963a). The materials consist 
of the same types of sediments as those that constitute the Haw- 
thorne Formation, that is, various combinations of quartz sand, 
clay, phosphate particles, and carbonate. In fact, it is only by 
geographic locality that some Bone Valley sediments can be dis- 
tinguished from Hawthorne materials. Consequently, the same 
problems are faced in naming and describing the Bone Valley 
sediments as are encountered in treating sediments of the Haw- 
thorne Formation. 

Loose sands usually ranging in thickness from a few feet to 
more than 25 feet are present as a surface blanket throughout the 
Bone Valley district. These sands rest upon underlying Bone Val- 
ley sediments. The Bone Valley materials often can be divided 
into two zones, an upper clayey sand to sand zone commonly 5 to 
10 feet in thickness, and a lower phosphorite zone commonly 15 



20 Quarterly Journal of the Florida Academy of Sciences 

to 25 feet in thickness (Sellards, 1910, p. 33; Cathcart, 1963a, b, c). 
These phosphatic Bone Valley sediments in turn rest throughout 
much of the district upon impure limestones, marls, and other 
sediments of the main body of the Miocene Hawthorne Formation. 
According to Cathcart (1963a, b, c), the commercial phosphate pro- 
duction comes from the lower phosphorite zone of the Bone Valley 
sediments and from the upper part of the Hawthorne materials. 
Details in distribution and thicknesses of the various rock ma- 
terials in the region are given by Cathcart (1950; 1963a, b, c). 

TABLE 7 

Peace River phosphate mine, Polk County, Florida (Pirkle et al., 1964) 
Surface sands and phosphatic sediments of the Bone Valley District * 







Insoluble residue 










Quartz 


Clay 


Total 


Total 






sand 


(—325 mesh) 


soluble 


p 2 o 5 


heavies 


Sample 


Unit 


in % 


in % 


in % 


in % 


in % 


1 


10 


98.10 


1.50 


.40 


.62 


.38 


2 


10 


98.30 


1.60 


.10 


.30 


.59 


3 


10 


91.18 


3.27 


5.55 


.32 


.39 


4 


9 


47.92 


35.80 


16.15 


11.08 


.12 


5 


9 


55.12 


20.40 


24.06 


10.63 


.08 


6 


9 


59.48 


21.29 


18.89 


9.34 


.07 


7 


9 


48.32 


14.77 


36.73 


13.42 


.07 


8 


8 


42.77 


13.77 


43.27 


13.30 


.10 


9 


8 


36.79 


13.17 


49.71 


17.60 


.11 


10 


8 


43.09 


24.06 


32.75 


9.12 


.10 


11 


7 


15.20 


19.24 


65.36 


25.23 


.04 


12 


7 


29,34 


20.25 


50.19 


26.14 


.08 


13 


7 


8.28 


18.48 


72.84 


16.08 


.03 


14 


7 


23.68 


17.03 


59.06 


27.26 


.06 


15 


6 


12.76 


21.44 


65.72 


21.65 


.05 


16 


6 


56.47 


15.55 


27.11 


9.79 


.33 


17 


6 


40.34 


10.35 


49.11 


16.23 


.17 


18 


5 


35.65 


11.96 


52.25 


16.13 


.13 


19 


4 


46.66 


8.20 


44.96 


17.79 


.30 


20 


3 


46.17 


6.78 


46.87 


20.53 


.25 


21 


2 


54.51 


10.24 


35.11 


11.39 


.20 


22 


1 


6.94 


19.85 


73.21 


2.33 


.03 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 21 

Peace River Mine 

Approximately 35 feet of sediments are exposed in mining op- 
erations at the Peace River phosphate mine (see section in Ap- 
pendix). There a relatively thin blanket of loose surface sands 
(unit 10 of Table 7) overlies 20 to 25 feet of Bone Valley sedi- 
ments. At the bottom of the mine pale yellow bedrock (unit 1) 
of the Hawthorne Formation is exposed. The features well shown 
in faces at this mine include the lenticular nature of the Bone Val- 
ley materials; the horizontal, laminated nature of the upper part 
of the phosphorite zone; and the lack of recognizable bedding in 
the lower part of the phosphorite zone just over the pale yellow 
bedrock. In addition, contact relationships between the pale yel- 
low bedrock and overlying phosphatic sediments are well shown. 
Likewise, the contact between the phosphorite zone of the Bone 
Valley materials and the overlying leached, vesicular sandrock is 
exceptionally clear. Tables 7-9 illustrate characteristics of the 
beds. 

Two well-developed organic zones are present, one at the base 
of the loose surface sands (unit 10) and the other near the base of 
the more highly vesicular sandrock of the leached zone (unit 9). 
Such organic zones may develop within sediments long after sedi- 
ments accumulate and thus cannot be considered, without some 
type of definite evidence, to represent soil zones or organic zones 
formed before overlying sediments were laid down. 

As seen from Table 9 the Bone Valley materials, like those of 
the main body of the Hawthorne Formation, are characterized by 
the presence of garnet. In general the total heavy mineral con- 
tent (excluding phosphorite and mica) is correlative with the con- 
tent of clastic quartz sand in the units, that is, the lower the quartz 
sand content the lower the percentage of heavy minerals (samples 
11, 13, and 22, Table 7 and Table 10). In some cases the percent- 
ages of heavy minerals can be shown to be further related to grain 
size of the quartz sand and to the degree of sorting of the sedi- 
ments. 

It is of interest to note that the heavy minerals may give in- 
sights into possible conditions that existed in the Piedmont-Blue 
Ridge region during the time that vast quantities of phosphatic 
materials were accumulating in Hawthorne and Bone Valley seas. 
The Piedmont-Blue Ridge area, generally considered to be the 



22 Quarterly Journal of the Florida Academy of Sciences 

original source area of the clastic quartz sand and the metamorphic- 
igneous heavy mineral suites of the Hawthorne and Bone Valley 
sediments, was furnishing large amounts of relatively easily weath- 
erable garnet to the seas. Release and dispersal of large amounts 
of garnet suggest that the source area was being eroded too rap- 
idly for the garnet to be destroyed through weathering, or else that 
the climate there was not a humid sub-tropical one as it is today. 



TABLE 8 



Peace River phosphate mine, Polk County, Florida 
Mechanical analyses of quartz sand fraction of various units (Pirkle et al. 



1964)* 







Quartz 


















sand 


Per 


cent total quartz 


sand retained on 


mesh 


Sample 


Unit 


in % 


10 


18 


35 


60 


120 


230 


1 


10 


98.10 


0.00 


,38 


5.42 


33.74 


34.30 


26.16 


2 


10 


98.30 


0.00 


.25 


5.90 


36.27 


33.99 


23.59 


3 


10 


91.18 


0.00 


.43 


5.79 


32.98 


34.69 


26.10 


4 


9 


47.92 


0.00 


.29 


9.40 


45.42 


40.56 


4.33 


6 


9 


59.48 


0.00 


.44 


11.06 


38.58 


44.76 


5.17 


7 


9 


48,32 


0.00 


.41 


6.48 


31.72 


56.42 


4.96 


9 


8 


36.79 


0.00 


,39 


5.72 


38.68 


43.02 


12.21 


10 


8 


43.09 


0.00 


.09 


9.76 


43.93 


41.75 


4.46 


12 


7 


29,34 


0.00 


.27 


12.87 


28.05 


48.85 


9.96 


13 


7 


8.28 


0.00 


trace 


16.59 


47.36 


24.52 


11.54 


14 


7 


23.68 


0.00 


trace 


17.23 


66.81 


11.26 


4.71 


15 


6 


12.76 


0.00 


trace 


18.44 


55.00 


21.72 


4.84 


16 


6 


56.47 


0.00 


trace 


1.27 


9.42 


28.73 


60.57 


17 


6 


40,34 


0.00 


.45 


10.39 


48.22 


36.45 


4.50 


18 


5 


35.65 


0.00 


trace 


5.77 


45.88 


45.66 


2.69 


19 


4 


46.66 


0.00 


.60 


5,34 


37.58 


51.18 


5.30 


20 


3 


46.17 


0.00 


.13 


7,30 


47.78 


38.75 


6.05 


21 


2 


54.51 


0.00 


,33 


12,34 


55.07 


27.66 


4.60 


22 


1 


6.94 


0.00 


.29 


6.61 


35.63 


52.87 


4.60 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix) and Table 7. Where available spot samples were used in 
preference to channel samples. 



Pirkle, Yoho, Allen: Analyses of Sediments 

TABLE 9 

Peace River phosphate mine, Polk County, Florida 
Heavy minerals in 1/16 to 1/8 mm fraction * 







CD 




CD 
C 
CD 




9£ 


CD 


CD 






o 




-M 




X 




CD '3 


•■-j 


13 


CD 






+J 


"3 

CD 


CD 


O 

o 


o 
o 


11 


o 
H 


g 


-4-> 

o 


"CD 

c 

J-c 


d 


3 


Jj 


"3 


CD 


£3 


>« 


a 


C 


a 




C/3 


£ 


5=1 


pc 


J 


N 


W "3 


00 


H 


w 


O 


1 


10 


A 


c 


A 


A 


c 


VR 


VR 








2 


10 


VC 


c 


A 


VC 


c 


R 


VR 


— 


— 


3 


10 


A 


c 


A 


A 


c 


R 


— 


— 


— 


4 


9 


VA 


R 


R 


C 


VC 


C 


R 


— 


VR 


5 


9 


A 


VR 


R 


C 


VC 


C 


R 


VR 


R 


6 


9 


A 


VR 


R 


C 


VC 


C 


R 


— 


R 


7 


9 


A 


VR 


R 


C 


VC 


C 


R 


VR 


R 


8 


8 


A 


R 


R 


C 


VC 


C 


R 


R 


R 


9 


8 


A 


R 


C 


C 


VC 


C 


R 


R 


R 


10 


8 


A 


R 


R 


VC 


VC 


C 


R 


R 


R 


11 


7 


A 


C 


R 


VC 


R 


C 


VR 


— 


R 


12 


7 


A 


R 


R 


VC 


c 


C 


R 


VR 


R 


13 


7 


VA 


C 


R 


VC 


R 


C 


VR 


VR 


R 


14 


7 


VA 


C 


R 


VC 


R 


C 


VR 


R 


R 


15 


6 


A 


c 


R 


VC 


R 


C 


— 


VR 


VR 


16 


6 


VA 


R 


C 


c 


C 


C 


— 


R 


VR 


17 


6 


A 


R 


C 


VC 


C 


C 


VR 


VR 


R 


18 


5 


A 


C 


R 


VC 


C 


C 


— 


R 


C 


19 


4 


A 


c 


C 


VC 


C 


C 


VR 


R 


R 


20 


3 


A 


R 


R 


c 


VC 


C 


VR 


C 


R 


21 


2 


A 


R 


R 


c 


VC 


C 


R 


c 


R 


22 


1 


A 


C 


C 


VC 


C 


C 


VR 


R 


R 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 

Achan Phosphate Mine 

At the site of the section measured at the Achan phosphate mine 
approximately 35 feet of phosphatic sediments are exposed (see 
Appendix). The phosphatic sediments are overlain by a blanket 
of loose surface sands (unit 10). Pale yellow bedrock (unit 1) is 
exposed at the bottom of the pebble mine. The phosphatic ma- 
terials exposed in the mine face clearly are divisible into an upper 
sand zone (units 6-9) and a lower phosphorite zone (units 2-5). 



24 Quarterly Journal of the Florida Academy of Scdznces 

Again garnet is found to characterize the Bone Valley sediments. 
Features of the materials are illustrated by data in Tables 10-12. 
Beautifully developed trough or scour cross-bedding was ob- 
served within the Bone Valley matrix only a few hundred feet 

TABLE 10 

Achan phosphate mine, Polk County, Florida 
Surface sands and phosphatic sediments of the Bone Valley district * 







Insoluble residue 


Total 








Quartz 


Clay 


Total 






sand 


(—325 mesh) 


soluble 


p,o 5 


heavies 


Sample 


Unit 


in % 


in % 


in % 


in % 


in % 


1 


10 


94.60 


1.81 





.48 


.24 


2 


10 


93.16 


2.20 


1.00 


.40 


.21 


3 


10 


88.27 


4.80 


3.20 


.38 


.27 


4 


9 


85.22 


13.79 


.90 


1.25 


.27 


5 


9 


78.30 


19.57 


1.99 


2.82 


.25 


6 


9 


82.60 


14.74 


1.80 


2.51 


.28 


7 


9 


88.38 


10.66 


.80 


1.71 


.22 


8 


8 


89.89 


7.12 


1.59 


1.81 


.30 


9 


8 


78.41 


15.57 


5.48 


2.39 


.32 


10 


7 


34.65 


53.59 


9.18 


4.56 


.14 


11 


6 


86.32 


8.51 


4.71 


1.87 


.58 


12 


5 


28.23 


9.92 


61.61 


17.67 


.10 


13 


5 


39.87 


18.69 


41.08 


11.15 


.19 


14 


5 


32.46 


6.31 


60.83 


18.55 


.13 


15 


4 


15.39 


6.03 


78.44 


26.39 


.06 


16 


4 


30.21 


12.89 


56.66 


16.66 


.13 


17 


4 


27.70 


9.59 


62.63 


20.46 


.10 


18 


3 


56.90 


9.92 


33.12 


10.62 


.14 


19 


3 


60.90 


17.62 


21.40 


6.84 


.24 


20 


3 


62.14 


20.59 


17.11 


5.20 


.22 


21 


2 


29.15 


24.27 


46.49 


13.29 


.10 


22 


2 


35.29 


17.37 


46.98 


11.54 


.09 


23 


2 


16.35 


13.71 


69.86 


21.38 


.07 


24 


2 


22.73 


8.69 


68.53 


19.51 


.11 


25 


2 


23.52 


8.92 


67.48 


21.70 


.10 


26 


2 


19.12 


15.38 


65.43 


13.27 


.06 


27 


1 


6.93 


4.59 


88.42 


6.82 


.03 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 25 

from the locality at which the section was measured. Such occur- 
rences of cross-bedding within the Bone Valley matrix are primary 
depositional features and as such, along with other primary de- 
positional features, constitute evidence that must be considered 
when studying possible origins for the sediments. Readings were 
made in 7 different sets of the cross-laminations. The mean direc- 

TABLE 11 

Achan phosphate mine, Polk County, Florida 
Mechanical analyses of quartz sand fraction of various units * 







Quartz 


















sand 


Per 


cent total quartz 


sand retained on 


mesh 


Sample 


Unit 


in % 


10 


18 


35 


60 


120 


230 


1 


10 


94.60 




.08 


1.82 


19.46 


35.33 


43.52 


2 


10 


93.16 




.09 


2.10 


21.31 


35.82 


40.69 


3 


10 


88.27 




.14 


1.77 


17.91 


33.80 


46.38 


4 


9 


85.22 




.09 


1.26 


38.46 


58.59 


1.59 


5 


9 


78.30 




1.30 


9.75 


50.10 


37.02 


1.83 


6 


9 


82.60 


.07 


1.29 


15.27 


44.76 


25.70 


12.90 


7 


9 


88.38 


.07 


.43 


6.20 


48.94 


42.49 


1.87 


8 


8 


89.89 




.53 


9.56 


47.38 


27.67 


14.85 


9 


8 


78.41 




.86 


8.85 


41.54 


41.64 


7.12 


10 


7 


34.65 


.17 


.81 


8.58 


41.99 


31.74 


16.71 


11 


6 


86.32 




.58 


5.45 


20.21 


61.69 


12.06 


12 


5 


28.23 


.98 


3.70 


24.44 


38.55 


28.56 


3.77 


13 


5 


39.87 




2.15 


17.16 


39.57 


35.92 


5.20 


14 


5 


32.46 




1.91 


20.74 


43.45 


29.66 


4.25 


15 


4 


15.39 




.26 


5.97 


43.97 


42.67 


7.13 


16 


4 


30.21 




.26 


4.29 


31.18 


54.16 


10.11 


17 


4 


27.70 




.22 


4.33 


35.69 


50.76 


9.01 


18 


3 


56.90 




.11 


2.77 


39.52 


52.91 


4.70 


19 


3 


60.90 






2.33 


33.66 


53.60 


10.41 


20 


3 


62.14 




.10 


.84 


21.42 


66.96 


10.69 


21 


2 


29.15 




.20 


4.58 


33.95 


52.66 


8.61 


22 


2 


35.29 




.17 


3.62 


38.16 


50.08 


7.97 


23 


2 


16.35 




.49 


4.64 


33.58 


49.33 


11.97 


24 


2 


22.73 




.44 


1.93 


17.35 


68.01 


12.27 


25 


2 


23.52 




.34 


4.16 


25.21 


54.33 


15.96 


26 


2 


19.12 




.42 


3.13 


32.08 


57.05 


7.31 


27 


1 


6.93 




.58 


5.76 


31.41 


44.96 


17.29 



Sample and unit numbers correspond to those of the measured section 



(see Appendix). 



26 Quarterly Journal of the Florida Academy of Sciences 



TABLE 12 

Achan phosphate mine, Polk County, Florida 
Heavy minerals in 1/16 to 1/8 mm fraction * 



C/3 


'2 


"2 




CD 
C 
1) 
X 

o 
o 

_1 


c 
o 
o 

•— 


"0 

1.1 


S-i 

5a 


<d 
c 

Eh 

C 
Eh 


CD 

w 




1 


10 


A 


C 


A 


VC 


c 


VR 


R 








2 


10 


VC 


C 


A 


c 


c 


R 


R 


— 


— 


3 


10 


VC 


c 


A 


c 


c 


R 


R 


— 


— 


4 


9 


VA 


R 


R 


c 


c 


C 


R 


— 


— 


5 


9 


VA 


R 


R 


c 


c 


C 


R 


— 


VR 


6 


9 


A 


R 


R 


c 


c 


C 


R 


— 


— 


7 


9 


A 


R 


R 


c 


c 


C 


R 


— 


— 


8 


8 


A 


VR 


C 


VC 


c 


R 


R 


— 


VR 


9 


8 


A 


R 


VR 


A 


c 


R 


VR 


VR 


R 


10 


7 


A 


R 


VR 


A 


c 


R 


VR 


VR 


VR 


11 


6 


A 


R 


R 


A 


c 


C 


R 


VR 


R 


12 


5 


A 


R 


R 


A 


c 


C 


VR 


C 


R 


13 


5 


A 


R 


C 


A 


c 


C 


VR 


C 


R 


14 


5 


A 


R 


R 


VC 


c 


R 


— 


C 


R 


15 


4 


A 


VR 


R 


c 


c 


C 


VR 


VC 


R 


16 


4 


A 


R 


R 


VC 


c 


C 


VR 


VC 


C 


17 


4 


A 


R 


R 


c 


c 


C 


— 


VC 


R 


18 


3 


A 


R 


R 


c 


c 


C 


VR 


A 


C 


19 


3 


A 


R 


R 


R 


VC 


C 


VR 


VC 


C 


20 


3 


A 


R 


R 


R 


VC 


C 


R 


VC 


R 


21 


2 


A 


— 


R 


C 


c 


C 


— 


C 


R 


22 


2 


A 


R 


R 


C 


c 


R 


R 


C 


C 


23 


2 


VA 


R 


R 


C 


c 


R 


R 


R 


C 


24 


2 


A 


R 


R 


C 


c 


C 


— 


R 


C 


25 


2 


A 


VR 


R 


R 


VC 


C 


R 


R 


C 


26 


2 


A 


R 


C 


C 


c 


C 


VR 


R 


R 


27 


1 


A 


R 


R 


C 


c 


C 


R 


VR 


R 



* Sample and unit numbers correspond to those of the measured section 
(see Appendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 27 

lion of greatest dip of the cross-laminations was determined to be 
S 44 degrees W. The average value for the maximum dip was 
calculated as 15 degrees. Attempts have not been made to deter- 
mine correlations between cross-bedding directions and such fea- 
tures as the thickness and grade of the matrix, but such studies 
should prove of unusual interest. 

In review, the most common heavy minerals of the Hawthorne 
Formation (Devil's Mill Hopper and Brooks Sink sections) and the 
phosphatic sediments of the Bone Valley district (Peace River and 
Achan sections) are ilmenite, zircon, kyanite, garnet, epidote, staur- 
olite, rutile, leucoxene, tourmaline, and sillimanite. These sedi- 
ments are characterized by the presence of persistent garnet and 
locally by relatively large amounts of epidote. Isphording (1963, 
pp. 19-22) has given petrographic descriptions of the various types 
of heavy minerals from the Hawthorne Formation. Hunter (1949, 
p. 415) has given descriptions of various kinds of heavy minerals 
from Bone Valley materials. 

CiTRONELLE FORMATION 

The Citronelle Formation in peninsular Florida (Cooke, geo- 
logic map, 1945) consists of quartz sands and clayey sands, locally 
containing important percentages of quartz or quartzite granules 
and quartzite pebbles. In some of the sediments white mica is a 
conspicuous constituent. The dominant clay mineral is kaolinite. 
This is present as a binder between sand grains and occasionally 
as stringers and small lenses of more nearly pure clay. Many of 
the quartzite pebbles are discoid; some are oval. The largest di- 
ameter of individual pebbles usually is less than IV2 inches, al- 
though some pebbles with lengths slightly more than 3 inches have 
been collected. 

The Citronelle sediments do not present any particular prob- 
lem in classification and can be given specific names when me- 
chanical analyses are available. The procedure used in this re- 
port for naming and describing the sediments has been described 
elsewhere (Pirkle and Yoho, 1961, pp. 252-253). 

Clermont Sand Mine 

The best exposures of Citronelle sediments in Lake County are 
at the Clermont sand mine of the Davenport Sand Company about 



28 



Quarterly Journal of the Florida Academy of Sciences 



4 miles east of Clermont in southern Lake County. There as much 
as 45 feet of these sediments have been exposed through mining 
operations. A section measured at this mine is given in the Ap- 
pendix. Features of the materials are illustrated in Tables 13-15. 



TABLE 13 

Clermont sand mine, Lake County, Florida 
Surface sands and Citronelle sediments (Pirkle et al., 1964)* 





Median 




Clay % 












(diam. 


Sorting 


(-325 


230 


120 


Heavies 


Heavies 


Unit 


mm) 


Coef. 


mesh) 


in% 


in% 


% on 230 


total % 


25 


.31 


1.35 


1.30 


2.08 


33.98 


11.42 


.35 


24 


.24 


1.95 


16.94 


4.74 


28.67 


3.22 


.26 


23 


.53 


2.01 


13.05 


1.69 


12.39 


1.89 


.07 


22 


.44 


1.91 


11.36 


3.85 


17.09 


1.48 


.16 


21 


.38 


2.20 


8.28 


10.20 


22.45 


1.56 


.20 


20 


.62 


1.84 


6.78 


3.08 


12.53 


1.94 


.13 


19 


.58 


1.50 


4.49 


3.88 


8.60 


2.34 


.22 


18 


.14 


1.85 


13.27 


28.38 


30.29 


1.72 


.50 


17 


.28 


2.98 


9.65 


18.07 


20.81 


1.30 


.35 


16 


.17 


2.19 


13.63 


19.16 


34.52 


1.43 


.35 


15 


.95 


1.39 


2.50 


1.91 


3.53 


1.18 


.11 


14 


.19 


1.73 


11.26 


16.37 


41.02 


1.64 


.36 


13 


.21 


1.46 


5.66 


7.71 


51.63 


3.43 


.28 


12 


.68 


1.67 


2.40 


2.31 


14.14 


4.02 


.13 


11 


.20 


1.65 


7.11 


12.17 


46.41 


4.29 


.48 


10 


.49 


1.95 


3.00 


2.08 


19.10 


4.13 


.10 


9 


.60 


2.00 


6.20 


7.60 


14.85 


1.90 


.13 


8 


.19 


2.02 


10.26 


17.16 


41.41 


1.73 


.34 


7 


.48 


1.78 


3.80 


3.16 


22.26 


4.65 


.25 


6 


.18 


1.33 


11.60 


11.47 


60.45 


2.27 


.33 


5 


.65 


1.81 


3.69 


1.92 


18.89 


4.06 


.07 


4 


.19 


1.20 


10.29 


4.66 


64.97 


5.70 


.34 


3 


.20 


1.25 


9.60 


3.32 


62.15 


7.17 


.39 


2 


.31 


1.70 


5.50 


2.87 


34.98 


4.46 


.14 


1 


Covered 
slope 















* Unit numbers correspond to those of the measured section (see Ap- 
pendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 



29 



Relationships of the loose surface sands to the surface topogra- 
phy and to the underlying clayey sands are exceptionally well 
shown at this locality. The base of the surface sands as seen in 
the pit faces is very irregular, with the surface sands at places pro- 
jecting abruptly downward for more than 40 feet as if filling old 
valleys or sinks. The upper surface of the sands forms a level to 
slightly undulating plain. It does not reflect topographically the 
old valleys or sinks. 



TABLE 14 

Clermont sand mine, Lake County, Florida 
Per cent by weight of various grade sizes in mm * 



Unit 


8-4 


4-2 


2-1 


i-% 


Vz-Va 


V4-V8 


y 8 -l/16 


<1/16 


25 




.05 


.62 


9.57 


52.15 


33.98 


2.08 


1.56 


24 


.82 


1.71 


8.23 


15.70 


22.77 


28.67 


4.74 


17.36 


23 




.77 


19.34 


32.08 


20.39 


12.39 


1.69 


13.33 


22 


.11 


.54 


10.46 


31.35 


24.90 


17.09 


3.85 


11.70 


21 




1.92 


13.46 


26.07 


17.25 


22.45 


10.20 


8.67 


20 




1.08 


23.60 


33.53 


19.20 


12.53 


3.08 


6.97 


19 


.37 


.66 


10.01 


46.95 


24.75 


8.60 


3.88 


4.79 


18 


.36 


.73 


7.07 


12.65 


6.87 


30.29 


28.38 


13.66 


17 


.20 


3.81 


19.99 


20.95 


6.03 


20.81 


18.07 


10.13 


16 


.23 


1.51 


9.75 


12.64 


8.26 


34.52 


19.16 


13.91 


15 


.20 


3.41 


41.10 


42.32 


4.85 


3.53 


1.91 


2.67 


14 




.50 


7.33 


11.35 


11.92 


41.02 


16.37 


11.52 


13 




.06 


.89 


8.86 


24.77 


51.63 


7.71 


6.08 


12 




.36 


21.19 


42.37 


17.02 


14.14 


2.31 


2.61 


11 




.15 


7.09 


13.43 


13.29 


46.41 


12.17 


7.47 


10 




.98 


21.36 


26.46 


26.80 


19.10 


2.08 


3.21 


9 




.68 


14.45 


39.59 


16.43 


14.85 


7.60 


6.40 


8 






7.15 


17.19 


6.59 


41.41 


17.16 


10.52 


7 




.09 


7.40 


40.33 


22.73 


22.26 


3.16 


4.03 


6 






3.28 


6.30 


6.53 


60.45 


11.47 


11.97 


5 




.10 


12.73 


48.45 


14.05 


18.89 


1.92 


3.85 


4 






1.21 


5.74 


12.98 


64.97 


4.66 


10.44 


3 




.03 


1.61 


6.80 


16.30 


62.15 


3.32 


9.78 


2 




.54 


5.96 


20.64 


29.31 


34.98 


2.87 


5.70 


1 


Covered slope 















* Unit numbers correspond to those of the measured section (see Ap- 
pendix). 



30 Quarterly Journal of the Florida Academy of Sciences 

Other features revealed in faces at this mine include excellent 
exposures of various combinations of thinly laminated sediments 
(units 5, 8, 16-18, 21, Table 14) that upon mechanical analyses give 
a bimodal distribution of the sand-size materials (usually low in 
60 mesh). The sands of the individual laminations are unimodal. 

TABLE 15 

Clermont sand mine, Lake County, Florida 
Heavy minerals in 1/16 to 1/8 mm fraction * 













"0 
















QJ 




9s 


V 


CD 








£ 




9 




03 C 


•s 


1i 


0) 




+j 


| 


jj 


6 
o 


c 
o 




o 
u 

2 


C 


o 

H3 


c 


£h 


£2 


3 


o 


(h 


>>s 


a 


o 


'3 




D 


^ 


~ 


J 


N 


WS8 


isi 


t_ 


w 


6 


25 


VA 


C 


R 


C 


R 


R 


VR 








24 


VA 


C 


R 


c 


VC 


R 


R 


— 


— 


23 


VA 


c 


R 


c 


VC 


R 


R 


— 


— 


22 


A 


R 


R 


R 


A 


C 


R 


— 


— 


21 


A 


C 


R 


R 


VC 


R 


R 


— 


— 


20 


VA 


R 


C 


C 


c 


R 


R 


— 


— 


19 


VA 


R 


R 


vc 


c 


R 


VR 


— 


— 


18 


VA 


C 


R 


c 


vc 


C 


R 


— 


— 


17 


VA 


R 


C 


c 


vc 


R 


R 


— 


— 


16 


A 


R 


C 


R 


vc 


C 


R 


— 


— 


15 


VA 


R 


C 


C 


vc 


R 


R 


— 


— 


14 


A 


R 


C 


C 


vc 


R 


R 


— 


— 


13 


VA 


R 


C 


C 


c 


R 


R 


— 


— 


12 


VA 


C 


R 


VC 


c 


R 


R 


— 


— 


11 


VA 


R 


C 


c 


c 


C 


R 


— 


— 


10 


VA 


R 


C 


c 


c 


R 


R 


— 


— 


9 


A 


C 


C 


c 


c 


C 


R 


VR 


— 


8 


A 


R 


C 


c 


vc 


R 


R 


— 


— 


7 


VA 


R 


C 


vc 


c 


R 


R 


— 


— 


6 


A 


C 


C 


R 


vc 


C 


R 


— 


— 


5 


VA 


R 


C 


c 


c 


R 


R 


— 


— 


4 


VA 


C 


C 


c 


vc 


C 


R 


— 


— 


3 


A 


C 


C 


c 


vc 


C 


R 


VR 


— 


2 


VA 


C 


R 


vc 


vc 


R 


R 


— 


— 


1 


Covered slope 














- - 



* Unit numbers correspond to those of the measured section (see Ap : 
pendix). 



Pirkle, Yoho, Allen: Analyses of Sediments 31 



TABLE 16 

Diamond Interlachen sand mine, Putnam County, Florida 
Heavy minerals in 1/16 to 1/8 mm fraction * 



"2 


£ 

'a 

0) 


1 


o 

8 
O 

i 


a 
o 
o 

H 

N 


H3 

«.§ 

<D 2 
CO J 


CO 

5a 


c 

o 


Epidote 
Garnet 


29 


VA 


c 


c 


C 


c 


c 


R 





28 


VA 


R 


R 


C 


c 


R 


— 


— — 


27 


VA 


R 


R 


C 


c 


R 


VR 


— — 


26 


A 


R 


R 


R 


vc 


C 


VR 


VR — 


25 


VA 


C 


R 


C 


c 


c 


— 


VR — 


24 


VA 


C 


VR 


C 


c 


R 


VR 


VR — 


23 


VA 


C 


C 


R 


vc 


C 


R 


VR — 


22 


VA 


C 


R 


R 


c 


C 


— 


VR — 


21 


VA 


C 


R 


C 


c 


C 


R 


— — 


20 


VA 


R 


R 


C 


c 


R 


VR 


— — 


19 


VA 


C 


R 


C 


R 


R 


VR 


VR — 


18 


VA 


c 


R 


R 


c 


R 


VR 


— — 


17 


VA 


R 


R 


R 


c 


C 


VR 


VR — 


16 


VA 


R 


R 


R 


vc 


C 


R 


— — 


15 


VA 


R 


R 


R 


vc 


C 


C 


— — 


14 


VA 


C 


C 


C 


c 


C 


VR 


— — 


13 


VA 


C 


R 


C 


c 


R 


VR 


— — 


12 


VA 


C 


R 


R 


c 


C 


R 


— — 


11 


VA 


R 


R 


C 


c 


C 


VR 


— — 


10 


VA 


C 


C 


R 


c 


C 


R 


— — 


9 


VA 


R 


R 


R 


vc 


R 


— 


VR — 


8 


A 


R 


R 


R 


A 


C 


R 


— — 


7 


VA 


C 


C 


C 


vc 


R 


VR 


— — 


6 


A 


C 


C 


C 


c 


C 


R 


— — 


5 


VA 


C 


R 


C 


c 


R 


— 


— — 


4 


VA 


C 


C 


R 


c 


C 


VR 


— — 


3 


VA 


R 


R 


R 


c 


C 


VR 


— — 


2 


A 


C 


C 


R 


vc 


C 


R 


VR — 


1 


Covered slope 















* Unit numbers correspond to those of the measured section (Pirkle et al v 
1963). 



32 Quarterly Journal of the Florida Academy of Sciences 

Units 6-7 (Table 14) can be selected to illustrate how various com- 
binations can result in bimodal distributions. These two units are 
of about the same thickness; the sands of each are unimodal. How- 
ever, if a channel sample was taken across both units (or if the 
analyses of the two units were combined), the sands of the com- 
bined sediments would be bimodal, low in 60 mesh. 

The most common heavy minerals of the Citronelle Formation 
are ilmenite, kyanite, zircon, staurolite, rutile, leucoxene, tourma- 
line, and sillimanite (Tables 15-17). The heavy minerals contain 
essentially no garnet and only minute amounts of epidote. In re- 
spect to these two minerals the heavy mineral suite of the Citro- 
nelle Formation is markedly different from that of the Hawthorne 
Formation. 

Diamond Interlachen Sand Mine 

Approximately 65-70 feet of Citronelle sediments are exposed 
at the Diamond Interlachen sand mine located at Interlachen, 
Putnam County, Florida. Lithologic features and characteristics 
of the sediments at this mine have been described (Pirkle et al., 
1963). These features include zones developed through weather- 
ing, the relationships of clay content to grain size and to sorting, 
the possible downward migration of clay within the sediments to 
leave behind loose surface sands, and cross-bedding. Heavy min- 
eral data for this sand mine have not been published and are here 
given in Table 16. The units in this table correspond to those 
on the detailed section previously published (Pirkle et al., 1963, 
pp. 137-141). 

Grandin Sand Mine 

Approximately 65 feet of Citronelle sediments are exposed at 
the Grandin sand mine near Grandin in northern Putnam County. 
Among features that have been observed in faces at this mine are 
various zones developed through weathering, prominent trough 
or "scour" type cross-bedding, draping of sands and clayey sands 
as a result of solution in underlying soluble sediments, and brecci- 
ation associated with the draping. Sediments exposed at this site 
have been described (Pirkle et al., 1963) and a detailed section 
given. However, no heavy mineral data have been published for 
this mine. Table 17 gives these data. The units in the table cor- 
respond to those on the detailed section previously published 
(Pirkle et al., 1963, p. 142-149). 



TABLE 17 
Grandin sand mine, Putnam County, Florida 
Heavy minerals in 1/16 to 1/8 mm. fraction 



-t-» 


03 

•a 

CD 
J 


CD 


CD 
C 
CD 
X 

o 
o 

CD 


a 

o 
o 

H 

N 


T3 
§0> 

cd *a 

II 


CD 

'c 


CD 
C 

| 
o 

Eh 


0) 

o 

'a 
w 


1 

o 


41 


VA 


R 


c 


VC 


c 


R 


R 


— 


— 


40 


VA 


C 


R 


vc 


c 


R 


VR 


— 


— 


39 


VA 


C 


R 


VC 


R 


R 


VR 


— 


— 


38 


VA 


R 


R 


vc 


c 


R 


VR 


— 


— 


37 


VA 


C 


R 


c 


C 


R 


— 


— 


— 


36 


VA 


C 


R 


c 


C 


R 


VR 


— 


— 


35 


VA 


C 


C 


c 


C 


R 


R 


— 


— 


34 


VA 


C 


R 


c 


VC 


c 


VR 


— 


— 


33 


VA 


C 


C 


vc 


c 


R 


VR 


— 


— 


32 


VA 


C 


C 


c 


vc 


R 


R 


— 


— 


31 


VA 


C 


R 


c 


c 


R 


R 


— 


— 


30 


A 


R 


C 


R 


A 


C 


R 


— 


— 


29 


VA 


C 


C 


c 


C 


R 


R 


— 


— 


28 


VA 


C 


C 


vc 


C 


R 


VR 


— 


— 


27 


VA 


C 


C 


vc 


C 


C 


R 


— 


— 


26 


VA 


C 


c 


c 


C 


C 


R 


— 


— 


25 


A 


c 


c 


R 


VC 


C 


R 


VR 


— 


24 


A 


c 


R 


R 


vc 


C 


R 


— 


— 


23 


VA 


c 


c 


R 


vc 


C 


R 


— 


— 


22 


A 


c 


R 


R 


vc 


C 


R 


— 


— 


21 


VA 


c 


R 


VC 


c 


C 


R 


VR 


— 


20 


VA 


c 


C 


C 


c 


C 


R 


— 


— 


19 


A 


c 


R 


VC 


c 


C 


R 


— 


— 


18 


VA 


c 


C 


C 


c 


C 


R 


— 


— 


17 


VA 


R 


R 


C 


c 


C 


VR 


— 


— 


15 


A 


c 


C 


VC 


c 


C 


R 


VR 


— 


14 


VA 


c 


C 


C 


c 


C 


VR 


— 


— 


13 


VA 


R 


C 


C 


c 


C 


R 


— 





12 


VA 


C 


C 


VC 


c 


R 


R 


— 





11 


VA 


C 


C 


c 


vc 


R 


R 


— 





10 


VA 


R 


R 


c 


vc 


R 


R 


— 





9 


A 


R 


C 


c 


vc 


R 


R 


— 


— 


8 


VA 


C 


C 


c 


vc 


R 


VR 


— 





7 


A 


R 


C 


c 


A 


R 


R 


VR 





6 


A 


R 


C 


R 


A 


C 


R 


— 





5 


A 


C 


C 


C 


vc 


R 


R 


— 





4 


A 


R 


C 


R 


A 


C 


R 


— 





3 


VA 


C 


C 


VC 


VC 


R 


R 


— 





2 


A 


C 


C 


C 


vc 


R 


R 








1 


Covered slope 

















34 Quarterly Journal of the Florida Academy of Sciences 

Uses of Heavy Mineral Data 

Uses of the types of data given in this report are almost un- 
limited. Heavy mineral analyses can be singled out as an illustra- 
tion. Heavy mineral suites of various formations differ. Further- 
more, variations of heavy minerals may occur within the same for- 
mation. The differences can be utilized as a tool in attacking many 
types of geological problems, both regional and local. In order 
to be more specific in demonstrating the variety of uses of heavy 
minerals, their applications to present controversies of the Bone 
Valley district of Florida will be indicated. 

Surface Sands of Bone Valley District 

A blanket of loose surface sands is present over much of Flor- 
ida. These sands have different origins. In some areas the sands 
may represent a weathering residuum. In other areas the sands 
may be wind-blown. Large regions are covered by loose sands 
that may have been reworked by streams "meandering" down 
slope. Extensive areas of the sands are marine, the origin often 
being associated with changes in sea level during Pleistocene or 
other late Cenozoic epochs. As can be demonstrated by recent 
studies in the Bone Valley district, there is a current trend to 
reconsider previous origins and to conclude that more and more 
areas of the loose surface sands are a residual blanket developed 
in situ from weathering. 

To illustrate, Altschuler and Young (1960, p. B 202) in their 
recent rejection of the concept that the surface sands of the Bone 
Valley district are marine Pleistocene sediments and in their sup- 
port of a strictly weathering phenomenon for the origin of the 
sands state, "The contact between the sand mantle and the under- 
lying clayey sands of the Pliocene Bone Valley formation is irreg- 
ular and gradational in detail. Patches of clayey sand occur in the 
sand mantle above the 'contact' and nests of eluviated loose quartz 
sand occur in the clayey sand beneath it. Size analyses and heavy 
mineral analyses of closely spaced samples through several verti- 
cal sections reveal that the sand blanket is essentially identical to 
the sand fraction of the subjacent Pliocene Bone Valley formation 
(fig. 89.2). Where observed along extensive plains, the sand mantle 
thins with corresponding thinning in the zone of lateritically al- 
tered clayey sand beneath it. These facts indicate that the quartz 



Pirkle, Yoho, Allen: Analyses of Sediments 35 

sand blanket is mainly an insoluble residue of the lateritic altera- 
tion of the Bone Valley formation, and not a transgressive Pleisto- 
cene deposit." Altschuler and Young (1960, p. B 203) conclude, 
"... Lateritic weathering has created a residual sand plain over 
the region, which preserves the original sediment differentiation." 

Heavy mineral data and sedimentary features given in the 
present report make it difficult to consider the sand blanket at the 
site of the measured sections as a residual sand plain developed 
in situ through weathering. This does not deny that the surface 
sands may represent sands reworked from underlying sediments. 
An examination of the sand analyses given in Tables 8 and 11 
shows that at the locality where the sections were measured the 
size distribution of the quartz sands of the surface blanket is dif- 
ferent from the size distribution of the quartz sands in the under- 
lying Bone Valley sediments (note particularly the 230 mesh frac- 
tion). There is no material now present beneath the loose surface 
sands that could weather in situ to give a residual blanket of sand 
with the distribution of quartz grain sizes found in the surface 
sands. Actually the sedimentary features of the surface sands 
reflect primary deposition. 

The sands reflect some post-depositional weathering. This is 
shown by the high percentages of leucoxene (samples 1-3, Table 
9; samples 1-3, Table 12). Likewise, some garnet may have been 
weathered from the upper part of the Bone Valley Formation. 

However, most of the heavy mineral features result from orig- 
inal deposition. To illustrate, it is well known that as the heavy 
mineral percentages increase in sediments, the heavy minerals 
with higher specific gravities may become more concentrated at 
the expense of heavy minerals with lower specific gravities (Mar- 
tens, 1935). An examination of samples 1-3 of Table 7 shows that 
the surface sands at the site where the section was measured at 
the Peace River mine are higher in heavy mineral content than 
the underlying sediments. Likewise the surface sands are rela- 
tively high in the heaviest of the heavy minerals (ilmenite plus" leu- 
coxene, rutile, and zircon) and relatively low in the lightest heavies 
as seen from a check of kyanite and sillimanite (Table 9). These 
relationships reflect primary deposition, not weathering. 

In conclusion, it is clear that considerations of the grain size 
distributions of the quartz sand and features of heavy minerals 
have a direct bearing on questions concerning the origin of the 



36 Quarterly Journal of the Florida Academy of Sciences 

blanket of surface sands. The uses of heavy mineral suites are not 
limited to studies of surface sands. Actually the heavies are of 
considerable interest in regard to other problems as can be illus- 
trated by implications concerning the origin of the phosphatic 
Bone Valley sediments. 

Residual as Compared to Depositional Origin of 
Bone Valley Sediments 

The origin of the Bone Valley sediments has been discussed 
in the literature for years. There are two dominant concepts aptly 
termed the residual hypothesis and the depositional hypothesis. 
Cathcart (1950, pp. 86-87) states, "The residual hypothesis of origin 
of the phosphate deposits in its extreme form attempts to account 
for the entire sequence of unconsolidated materials overlying the 
limestone of the Hawthorn formation in the land-pebble district 
as the relatively insoluble residue of the Hawthorn that has been 
concentrated in place by weathering." According to Cathcart the 
depositional hypothesis, "... postulates erosion of the limestone 
of the Hawthorn formation at the end of Miocene time, deposition 
in Pliocene time of the lower unit of the Bone Valley formation 
in a marine offshore environment, subsequent erosion, and finally 
deposition in Pleistocene time of surficial terrace sands." After 
giving facts supporting both hypotheses Cathcart concludes, "Evi- 
dence is lacking on which to establish either hypothesis as a fact 
at any specific place because many field relations are subject to 
two or more interpretations." In recent studies Ketner and Mc- 
Greevy (1959) and Carr and Alverson (1959) tend to support the 
residual hypothesis. Cathcart (1963a,b,c) supports the deposi- 
tional hypothesis. 

The distribution of heavy minerals at the site where the sec- 
tion was measured at the Achan mine is given in Table 12. The 
Achan mine was selected for illustration because there the two 
zones of the Bone Valley sediments are clearly distinguishable. 
Of special interest to the problem of origin is the distribution of 
the relatively easily weatherable heavy minerals, garnet and 
epidote. 

Unit 10 (samples 1-3) is the blanket of loose surface sands. 
Those sands carry essentially no garnet or epidote and have been 
subjected to rather severe weathering as attested by the relatively 



Pirkle, Yoho, Allen: Analyses of Sediments 37 

large amounts of leucoxene and the relatively small amount of 
ilmenite. Units 9 through 6 constitute the upper sand zone of 
the Bone Valley sediments. Samples collected from this upper 
sand zone are labeled 4-11. This zone is approximately 10 feet 
thick and is considered as overburden in mining operations. An 
occasional grain of garnet is present in this zone (see sample 5). 
The first few grains of epidote appear in unit 8. Thus the upper 
sand zone contains only very rare epidote and only a very small 
amount of garnet. The essential lack of garnet and epidote in 
this zone probably reflects weathering as does the low phosphate 
content (Table 10) and possibly as do clay types (Altschuler et al., 
1956; Reves, 1960). 

The lower phosphorite zone of the Bone Valley materials are 
units 5 through 2. This zone is known as the matrix; it is the com- 
mercial phosphate zone. Samples collected from the zone are 
numbered 12-26. This phosphate zone is 27 feet thick. It con- 
tains garnet and epidote throughout. Within this zone the distri- 
bution of garnet is compatible with a weathering phenomenon. 
However, from Table 12 it is seen that epidote, likewise a rather 
easily weatherable heavy mineral, is more abundant higher in the 
matrix (units 5 through 3) than lower in the matrix (unit 2). Ac- 
cording to a strictly weathering phenomenon, the higher parts of 
the matrix should be the most weathered parts. Therefore, the 
distribution of epidote would be difficult to explain if the Bone 
Valley beds were a residue developed in situ from the weathering 
of the underlying main body of the Hawthorne sediments. Nor 
could leaching by laterally moving ground water within more 
permeable beds be used to explain the relationships. The laterally 
moving ground water would have been effective in weathering 
the garnet as well as in weathering the epidote. The garnet dis- 
tribution does not show this effect. 

To recapitulate, the heavy minerals are of considerable use in 
evaluating and determining the origin of the Bone Valley sedi- 
ments. In the case of the Achan mine the distribution of heavy 
minerals seems incompatible with the concept that parent Haw- 
thorne limestones or marls weathered in situ to result in the 
various overlying zones. Rather, the distribution of heavies would 
suggest that the Bone Valley sediments were deposited over the 
main body of the Hawthorne Formation, as evidenced from the 
distribution of epidote in the phosphorite zone, and loose surface 



38 Quarterly Journal of the Florida Academy of Sciences 

sands were later deposited as a separate unit over the Bone Valley 
sediments. The heavies likewise indicate post-depositional weath- 
ering for the sequence, as reflected by the relationships of ilmenite 
to leucoxene in the surface sands and in the essential absence of 
garnet and epidote in the upper sand zone of the Bone Valley 
sediments. 

Acknowledgments 

This work was made possible through funds furnished by the 
Graduate School of the University of Florida and by the Univer- 
sity of Florida Engineering and Industrial Experiment Station. 

Without the counsel, advice, and encouragement of Harold L. 
Knowles of the Department of Physical Sciences this work could 
not have been initiated and would not have been completed. That 
department and the Department of Civil Engineering made avail- 
able the laboratory space and equipment utilized in making most 
of the analyses presented. Sand and phosphate companies have 
extended many kindnesses. The help from each is sincerely ap- 
preciated. 

O. W. Babb of the Gainesville Equipment Company and Allen 
C. Edgar of the Edgar Minerals Corporation and the Mid-Florida 
Mining Company have visited exposures through various parts of 
the peninsula with the writers and have contributed immeasurably 
to the work. 

The writers are deeply indebted to Pierce Brodkorb of the 
University of Florida and to William D. Reves of the Florida Ge- 
ological Survey for their careful and painstaking review of the 
report. The talents of these men are reflected in a more accurate 
and readable work. Thanks are extended to Miss Rosa M. Ortiz 
for typing materials from script difficult to decipher. To all of 
the individuals and organizations aiding in various phases of the 
study the writers express their sincere gratitude. However, only 
the authors should be held responsible for the contents of the 
report. 

Summary 

In this report detailed sections with analyses are given of Haw- 
thorne, Bone Valley, and Citronelle sediments of Florida. Haw- 
thorne sections were measured at the Devil's Mill Hopper in 
Alachua County and at Brooks Sink in Bradford County. Bone 
Valley sections were measured at the Peace River and Achan 



Pirkle, Yoho, Allen: Analyses of Sediments 39 

mines in Polk County. A Citronelle section was measured at the 
Clermont sand mine in Lake County. Various analyses given in- 
clude the per cent quartz sand, mechanical analyses of the quartz 
sand, the per cent clay, the per cent soluble, the P 2 5 content of 
the sediments, the per cent heavy minerals, and heavy mineral 
analyses. Specific data showing the differences in the heavy min- 
eral suites of the Hawthorne and Citronelle formations of the 
peninsular are given for the first time in this report. 

Contrary to what has been suggested recently, the surface 
sand blanket within the Bone Valley district at the sites of the 
measured sections is not an insoluble residue developed in situ 
through weathering; the sands were laid down as a separate unit 
over the Bone Valley sediments. Furthermore, the lower phos- 
phorite zone and the upper sand or clayey sand zone of the Bone 
Valley are not a product from the weathering in situ of the under- 
lying Hawthorne Formation; the Bone Valley materials represent 
primary deposition of sediments. 

Appendix 

Devil's Mill Hopper 

Section 15, T. 9 S, R. 19 E, Alachua County, Florida 
(approximately 6 miles northwest of Gainesville)* 







Thickness 
(in feet 


Unit 


Description 


and 
inches) 



Surface Sands 

16. Sand. Loose, gray to white 3' 0' 

Sample 1. Spot sample of loose surface sands. 

Hawthorne Formation 

15. Phosphate concentration. Abundant pebbles and grains of phos- 
phorite embedded in a matrix consisting largely of quartz sand 
and clay. 
Many of the pebbles of phosphorite are impure limestone or marl 
fragments in which phosphate has replaced carbonate. These 
pebbles contain much included quartz sand. 
Most of the phosphate particles are some shade of white, gray, 

brown or black 24' 2" 

Sample 2. 1% foot channel sample of phosphatic sediments col- 
lected near top of unit. Southeast side of sink. Con- 
tains much pebble-size phosphorite. 



40 Quarterly Journal of the Florida Academy of Sciences 

Sample 3. Spot sample of phosphatic materials taken 5 feet 
down from upper surface of unit. Southeast side of 
sink. Contains much pebble-size phosphorite. 

Sample 4. Spot sample taken on west side of sink 16 feet down 
from upper surface of unit. Contains abundant fine 
phosphorite. 

14. Dolomitic limestone to dolomite. Cream to white to yellow, con- 
taining in places abundant molds and casts of marine pelecypods 
and gastropods. 
Locally quartz sand is an important constituent of the unit. In 

some places phosphate particles are common 11' 6" 

Sample 5. Spot sample of yellow dolomitic limestone or dolo- 
mite. 

13. Clayey sand. Yellow to yellow-brown with local occurrences of 
irregular masses of white carbonate. 
Upper 7 inches of unit has a greenish-blue color and a higher con- 
tent of quartz sand 3' 6" 

Sample 6. Spot sample of the yellow-brown clayey sand. 
Sample 7. Spot sample of the upper 7 inches of greenish-blue 

material. 
Sample 8. Spot sample of a nearly white carbonate mass. 

12. Clayey sand. Upper 1V2 feet of unit is dark blue; rest of unit is a 
light pastel greenish-blue. 
Pyritic T 2" 

Sample 9. Channel of all of unit. 

Sample 10. Spot sample of dark blue clayey sand near top of unit. 

Sample 11. Spot sample taken 2 feet 9 inches below upper surface 

of unit in light pastel, greenish-blue clayey sand. 
Sample 12. Spot sample of light pastel, greenish-blue clayey sand 

taken near base of unit. 

11. Conglomerate. Green to yellow. 

Unit consists of a mixture of quartz sand, clay and carbonate. 

The pebbles appear to be composed largely of quartz sand ce- 
mented with clay and/or carbonate. The pebbles break down 
readily when subjected to normal treatments used in making 
mechanical analyses. 

Locally black phosphate grains are common. 

Pyritic 2' 6" 

Sample 13. Spot sample taken near top of conglomerate. 

Sample 14. Spot sample taken near base of conglomerate. 

10. Calcareous clayey sand and sandy clay to massive, blocky clay. 

Light green to blue. 
Locally phosphate particles are common. In places the unit is 

highly calcareous. 
The clay present in this unit and continuing upward through unit 13 



Pirkle, Yoho, Allen: Analyses of Sediments 41 

has a different appearance from the clay of underlying units and 
usually has a darker color when fresh. 

Quartz sand, carbonate, and phosphate particles are more abundant 
in the upper 10 feet of the unit. Much of the lower 9 feet con- 
sists of massive, blocky clay with intercalated stringers and small 
lenses of sand and carbonate. Many of the clay blocks are sur- 
rounded by networks of quartz sand and carbonate. In the 
lower foot of the unit quartz sand increases. 

Because of slumpage some of the sediments in this interval could 
not be sampled or observed. Thus no attempt was made to sub- 
divide these sediments. 

Pyritic 20' 6" 

Sample 15. Spot sample taken near top of unit. 

Sample 16. Channel sample 2 feet and 2 inches in length, taken 
about 7 feet down from the top of the unit. 

Sample 17. Channel sample of the lower 4 feet 6 inches of unit. 

9. Massive clay. Gray to greenish-gray, blocky with networks of 
sand and carbonate surrounding some clay blocks and with string- 
ers of sand and carbonate within the clay 3' 6" 

Sample 18. Channel sample of all of unit. 

8. Clayey sand. Gray, soft I'll" 

Sample 19. Channel sample of complete unit. 

7. "Limestone." White to gray, lithified. Contains remnants of clay 
blocks. 

Forms nodular masses and slight ledges upon weathering 2' 0" 

Sample 20. A series of chips collected from all parts of unit. 

6. Clay. Gray to grayish-green to olive green, massive. 

Clay is blocky with networks of carbonate surrounding clay blocks 

and with stringers and small lenses of carbonate present within 

the clay. 
Carbonate apparently is replacing the clay. 
Analyses indicate that in some places the massive clay is calcareous 

and in other places non-calcareous 10' 4" 

Sample 21. Channel sample of all of unit. 

Sample 22. Spot sample of massive clay from unit. 

5. Zone containing intraformational breccias or conglomerates. Up- 
per part of unit is a prominent conglomerate. Near base of 
unit is another well-defined conglomerate. Several less conspic- 
uous conglomerates are present through the unit. 
The intraformational breccias or conglomerates consist of various 
mixtures of quartz sand, clay, phosphate particles, and carbonate. 
In places these zones contain calcite fossil shells, and angular 
blocks and rounded pebbles of grayish-green clay. The most 
prominent fossils are Pecten acanikos Gardner, Carolia floridana 

Dall, and barnacles 10' 11" 

Sample 23. Spot sample taken 1 foot 1 inch down from upper sur- 



42 Quarterly Journal of the Florida Academy of Sciences 

face of unit within the upper prominent conglomerate 

zone. 
Sample 24. Spot sample of blocky clay with networks of calcite 

and quartz sand. Taken 3 feet 6 inches below top 

of unit. 
Sample 25. Spot sample from a conglomerate zone 6 feet 9 inches 

below upper surface of unit. 
Sample 26. Spot sample taken V-k feet above base of unit. 
Sample 27. Spot sample taken 12 inches above base of unit in 

lower, well-defined conglomerate zone. 

4. Mixture of quartz sand, clay, and carbonate. Olive green. 

As indicated by Thompson and Floyd in an unpublished thesis 
by McClellan (1962), the distinctiveness of this unit is an aid in 

locating the units in the lower part of the Mill Hopper sink 1' 1" 

Sample 28. Channel sample of all of the olive green marker bed. 
Sample 29. Channel sample of all of unit. Collected approxi- 
mately l 1 /^ feet from site of sample 28. 

3. Clayey limestone. White, soft. Contains remnants of gray clay 

blocks, more numerous near upper part of unit 2' 5" 

Sample 30. Channel sample of all of unit. 

2. Covered slope. 

An occasional exposure. 

Approximately 6 1 /£ feet above base of unit calcareous sand is ex- 
posed. Just over unit 1, clay is exposed. Cannot be certain 
if these exposures represent materials in place or slumped sedi- 
ments 9' 1 " 

Sample 31. Spot sample of calcareous sand collected 6Y2 feet 
above base of unit. 

1. Sandy limestone. Locally dolomitic. White. Rests upon under- 
lying Ocala limestone. 

In places dense, dark colored, with stringers of quartz sand. Con- 
tains occasional blocks of gray clay. 

Parts of this unit are highly silicified. Where the silicified portions 

rest upon Ocala limestone, that limestone is often silicified 3' 2" 

Sample 32. Chips taken from all parts of the unit. 



Total depth 116' 9' 



* Measured by E. C. Pirkle, Fred Pirkle, and W. H. Yoho, early spring, 
1962. For analyses of samples see Tables 1-3. 



Pirkle, Yoho, Allen: Analyses of Sediments 43 

Brooks Sink 
Section 14, T. 7 S, R. 21 E, Bradford County, Florida* 

Thickness 
(in feet 
Unit Description and 

inches) 

20. Covered interval 

Although not visible at site of the section because of slumpage and 
vegetation, loose to slightly indurated surface sands blanket the 
region. Clay content increases with depth in these sands. In 
many areas lenses of nearly pure gray to greenish-gray clay are 
present near the base of this upper sand unit. 

The surface sands were sampled at a distance of approximately 

50 feet from the rim of the sink 15' 0" 

Sample 1. Spot sample of fine white surface sands. 

Sample 2. Spot sample of tan to brown sand taken 2 feet below 
land surface and directly beneath sample 1. 

Shell Marl (Late Miocene?) 

19. Shell marl, cream to light tan. 

Horizontal bedding distinct, some cross-bedding visible near top 
of unit. 

Quartz sand and clay increase with depth 20' 7" 

Sample 3. Spot sample taken in an oolitic zone 2 feet below 

upper surface of marl. 
Sample 4. Spot sample collected 10 feet down from top of unit. 
Sample 5. Spot sample taken 1 foot above base of unit. 

Sediments of Late Miocene age (?) (Possibly Hawthorne sediments). 

18. Quartz sand and finely divided carbonate containing shell frag- 
ments. Cream to tan. 
Numerous shiny black phosphate grains 4' 10" 

Sample 6. Spot sample taken 1 foot 2 inches below top of unit. 
Sample 7. Spot sample taken 3 feet below upper surface of 

unit. 
Sample 8. Channel sample of all of unit 18. 
17. Sandy limestone, clayey. Cream to light tan. 

Contains scattered grains of shiny black phosphorite. 

Occurs as a slightly developed ledge 1' 2" 

Sample 9. Representative sample of unit 17. 

Hawthorne Formation (Miocene) 

16. Mixture of carbonate, quartz sand and clay with common to abun- 
dant grains and pebbles of black and brown phosphorite. Tan 
to buff. 



44 Quarterly Journal of the Florida Academy of Sciences 

Fossil impressions of mollusks common. Occasional interior molds 

of large clams 1' 6" 

Sample 10. Sample representative of unit 16. 
15. Limestone, sandy. Tan to buff. 

Contains impressions of mollusks and a few grains of black, shiny 
phosphorite. 

Occurs as a slight ledge 10" 

Sample 11. Series of chips from all of unit 15. 
14. Mixture of carbonate, quartz sand, detrital limestone particles, and 
clay with common grains and pebbles of brown phosphorite. 
Tan to buff. 
Thin zone of limestone pebbles, some an inch in diameter, near 
base of unit. The limestone pebbles contain quartz sand, numer- 
ous pebbles and grains of phosphorite, and abundant impres- 
sions of fossils, both pelecypods and gastropods. Varying 
amounts of the carbonate content of these pebbles have been 

replaced by phosphate 1' 0" 

Sample 12. Limestone pebble partly replaced by phosphate, se- 
lected from thin zone of limestone pebbles near base 
of unit 14. 
Sample 13. Channel of all of unit 14. 
13. Limestone. Tan to buff. 

Contains common pebbles of brown phosphorite and impressions 
of mollusks. 

Occurs as a poorly developed ledge 6" 

Sample 14. Series of chips from all parts of unit 13. 
12. Mixture of carbonate, quartz sand, and clay with numerous peb- 
bles and grains of brown and black phosphorite. Tan to buff 
to pale yellow. 
Numerous impressions and molds of fossils, mainly pelecypods. 
Carbonate is more abundant in upper part of unit. 
Lower 8 inches of unit contains a heavy concentration of phos- 
phorite 2' 0" 

Sample 15. 1 foot 4 inch channel sample of the upper, more cal- 
careous part of unit 12. 
Sample 16. Channel sample of the lower 8 inch phosphate con- 
centration. 

11. Mixture of carbonate, quartz sand, and clay with common brown 
and black phosphate grains. Tan to buff to pale yellow. 
Impressions of fossil pelecypods 5' 0" 

Sample 17. Representative sample of unit 11. 

10. Limestone, sandy. Tan to buff. 

Contains common grains of black and brown phosphorite. 

Many fossil impressions, particularly pelecypods. 

Occurs as a ledge 1' 6" 

Sample 18. Chips from all of unit 10. 



Pirkle, Yoho, Allen: Analyses of Sediments 45 

9. Mixture of carbonate, quartz sand, and clay with common grains 
of black and brown phosphorite and numerous fossil impres- 
sions, mostly pelecypods. Gray. 

Weathers yellowish to tan and buff 4' 8" 

Sample 19. Spot sample collected approximately 2 feet below 

upper surface of unit 9. 
Sample 20. Spot sample collected near base of unit 9. 
Sample 21. Channel sample of all of unit 9. 
8. Sandy limestone, clayey. Tan to buff. 

Contains common brown and black phosphate grains. 

Occurs as a ledge 1' 0" 

Sample 22. Chips from all parts of unit 8. 
7. Mixture of quartz sand, carbonate, and clay with common to nu- 
merous grains of shiny black and brown phosphorite. Gray or 
blue-gray. 

Weathers tan or buff 3' 3" 

Sample 23. Spot sample of fresh, blue-gray material of unit 7. 
Sample 24. Channel sample of all of unit 7. 
6. Limestone, sandy and clayey. Gray or blue-gray. Contains small 
shiny black and brown phosphate grains. Weathers tan or buff. 

Occurs as a ledge 7' 5" 

Sample 25. A series of chips collected from all parts of unit 6. 
5. Mixture of quartz sand, carbonate, and clay. Gray or blue-gray. 
Contains numerous small black, brown, and gray phosphate grains. 
Weathers tan or buff. 
Scattered through unit are occasional white, poorly preserved 

pelecypods 2' 0" 

Sample 26. Sample representative of fresh, bluish material of 
unit 5. 
Diastem? 

4. Layer of broken-up, mollusk-bored, fine-grained limestone. Blue- 
gray. 
Weathers tan to buff. 

The limestone fragments and blocks vary from a fraction of an 
inch to more than 1 foot in longest dimension and are embedded 
in a matrix consisting of a mixture of carbonate, quartz sand, 
clay, and abundant pebbles and grains of black, brown, and gray 
phosphorite (Pirkle, Figures 7 and 8, 1956). 
Many white, poorly preserved fossil fragments of mollusks inter- 
mingled with and on top of the fragments of limestone 4" 

Sample 27. Sample of the matrix in which the broken fragments 

of the mollusk-bored limestone are embedded. 
Sample 28. Representative fragment of the mollusk-bored lime- 
stone. 
3. Phosphate concentration. Tan or buff colored. Blue where un- 
weathered. 



46 Quarterly Journal of the Florida Academy of Sciences 

Mixture of carbonate, quartz sand, and clay containing abundant 
pebbles and grains of phosphorite and rounded and angular 
blocks of clay. 

Pebbles of phosphorite are black, gray, and white. 

Grains of phosphorite are black, gray, and brown I'll" 

Sample 29. Channel of all of unit 3. 
2. Limestone, clayey. Cream or buff colored. 

Locally contains clusters of phosphate grains and pebbles. 

Bottom of unit has a wavy, very irregular surface and shows solu- 
tion weathering. 

Occurs as a ledge 1'10" 

Sample 30. Chips of the limestone ledge of unit 2. 
1. The upper one to two feet of this unit is a solution-riddled rock 
with the solution channels filled with a mixture of carbonate, 
quartz sand, clay, and abundant phosphate grains and pebbles. 
When this part of the unit is cleaned with a hammer the material 
has the appearance of a breccia with white carbonate fragments 
embedded in a matrix of carbonate, quartz sand, clay, and phos- 
phate particles (Pirkle, Figures 5 and 6, 1956). 

"Breccia" grades downward into blue-gray calcareous, more or 
less sandy, clay. This calcareous clay contains common pebbles 
and grains of phosphorite. Clay weathers tan or buff. 

Near bottom of stratum, in places, are numerous fragments of poor- 
ly preserved white pelecypod shells 5'10" 

Sample 31. Sample of material representative of unit beneath the 
"breccia." 

Sample 32. Sample of material occupying the solution channels 
in upper "breccia" part of unit 1. 
0. Mixture of carbonate, quartz sand, and clay. White to light tan 
or yellow. Blue where fresh. 

Contains scattered black, brown, and gray phosphate pebbles and 

grains 2' 0" 

Sample 33. Sample selected as representative of unit 0. 



Total depth to lake 84' 2" 

* Measured by E. C. Pirkle, W. H. Yoho, and O. W. Babb during April, 
1962. For analyses of samples see Tables 4-6. 



Pirkle, Yoho, Allen: Analyses of Sediments 47 

Peace River Phosphate Mine 

Section 12, T. 31 S., R. 25 E., Polk County, Florida, 
approximately 4Vz miles southeast of Bartow, Florida* 

Thickness 
(in feet 
Unit Description and 

inches) 

Surface Sands 

10. Surface sands, loose. Medium to very fine. Upper 3 feet 8 inches 
of unit is white to light gray. Lower 2 feet 7 inches of unit 
contains much organic matter and is dark brown to black. 
These sands are present as a blanket over the Bone Valley sedi- 
ments throughout area 6' 3" 

Sample 1. Spot sample taken near land surface in present root 

zone. 
Sample 2. Spot sample collected from a depth of approximately 

3 feet below land surface. 
Sample 3. Spot sample from the organic zone near base of unit. 

Phosphatic Sediments of Bone Valley 
9. Porous "sandrock." Leached zone. 

From top of unit to depth of 7 feet 4 inches sediments are white 
to tan to light brown. The materials are indurated as a result 
of weathering, and the phosphorite has been leached leaving 
a porous or vesicular sandstone or "sandrock." 
Beneath the upper leached white "sandrock" is a dark gray to black 
zone high in organic content (at times referred to locally as hard- 
pan). The "sandrock" of this dark zone also is vesicular; many 
of the original phosphate particles have been removed to leave 
the openings. 
The lower 1 foot 3 inches of the unit is not as black nor does it 
have as many openings as the overlying black organic zone. 
Also this lower zone has phosphate or soft clay in partially de- 
veloped openings. 

The quartz sand of this unit is fine to coarse 10' 6" 

Sample 4. Spot sample of whitish, vesicular "sandrock" selected 

approximately 6 feet below upper surface of unit. 
Sample 5. Channel sample of lower 3 feet 2 inches of unit (the 

black, vesicular "sandrock" and underlying non-vesicular 
sediment). 
Sample 6. Spot sample from the black, vesicular "sandrock." 
Sample 7. Spot sample from the basal, non-vesicular zone. 
8. Mixture of quartz sand, clay, and phosphate particles. Whitish 
throughout most of unit. Light bluish-gray to greenish-gray 
in basal 7 inches. 



48 Quarterly Journal of the Florida Academy of Sciences 

Phosphorite occurs as small pebbles and granules and as numerous 
sand-size particles. Most of the phosphorite is whitish to gray; 
some is brown. 

Quartz sand is medium to very fine 3'11" 

Sample 8. Channel sample of all of unit 8. 

Sample 9. Spot sample taken 1 foot down from upper surface 
of unit in the white material. 

Sample 10. Spot sample from the 7-inch greenish-gray basal zone. 
7. Mixture of clay, quartz sand, and phosphorite. 

Phosphate pebbles, granules, and sand-size particles are abundant. 

Upper 9 inches of unit is pale yellow to yellow-brown and likely 
represents a weathered zone of underlying sediments. In this 
zone there are occasional very large phosphate masses, some 
with diameter of 2 to 3 inches. 

In the middle 11 inches of this unit there is an increase in the 
quantity of pebble-size and granule-size phosphorite. The sedi- 
ments are whiter. 

The lower 10 inches of the unit is gray to lavender and is charac- 
terized by a marked increase in sand-size brown phosphorite. 

The phosphate pebbles in this unit are mostly white to gray to 
brown; the phosphate grains are white to brown to orange 
and amber. 

Quartz sand within the unit is coarse to very fine 2' 6" 

Sample 11. Channel sample of all of unit 7. 

Sample 12. Spot sample from upper, more weathered 9 inches 
of unit 7. 

Sample 13. Spot sample of the whiter, middle 11-inch zone of 
unit 7. 

Sample 14. Spot sample of lower 10-inch zone characterized by 
marked increase in sand-size, brown phosphorite. 
6. Mixture of quartz sand, clay, and phosphate particles. Yellow- 
brown to chocolate brown. 

Upper 9 inches of unit is a darker color (ranging from yellow- 
brown to drab to chocolate) and appears to be a decomposed 
and fragmented shell marl. The marl fragments contain many 
impressions of marine mollusks. The carbonate content of the 
masses of marl has been replaced to varying degrees by phos- 
phate. Most of the phosphorite of this zone is brown. The 
quartz sand is fine to coarse. 

The middle 20 inches of this unit contains a marked increase in 
quartz sand, 29 per cent of which is fine and 61 per cent of 
which is very fine. The color is yellow-brown. Pebble-size 
phosphorite has decreased sharply but sand-size white, gray, 
tan, brown to orange phosphorite is abundant. 

The basal 8 inches of unit is distinctly banded white. It is clear- 
ly defined and can be seen for tens of feet around the face of 
the mine. Quartz sand fine to coarse. Phosphate pebbles are 



Pirkle, Yoho, Allen: Analyses of Sediments 49 

not common but sand-size phosphate particles varying in color 
from white, to gray, to light brown, to orange are abundant. 

In regard to color, from unit 1 through unit 6 most sediments are 
some shade of yellow to yellow-brown with intercalated whitish 
lenses. Beginning with unit 7 and going upward, most of the 
pale yellow to yellow-brown colors are replaced by lavender, 
light gray to bluish, and greenish-gray to whitish colors 3' 1" 

Sample 15. Spot sample of upper 9-inch decomposed limestone 
or marl. 

Sample 16. Spot sample of middle 20-inch zone characterized by 
a sharp increase in very fine quartz sand. 

Sample 17. Spot sample of lower, white-banded 8-inch zone of 
unit 6. 
5. Mixture of quartz sand, clay, and phosphorite. Mottled yellow- 
brown and white. Lower part of unit is whiter in color. 

Horizontally laminated. Going upward from the bed rock of 
unit 1 this is the first unit in which horizontal laminations can 
be detected. Because of the laminations this would be the first 
unit of the Bone Valley Formation according to some investi- 
gators. Underlying units 4, 3 and 2 would be considered a 
weathering residuum and with unit 1 would be designated as 
Hawthorne sediment. 

Sand-size phosphorite is abundant. Only a minor quantity of peb- 
ble-size phosphate particles. The phosphorite is white, tan, 
brown, and orange. 

Quartz sand is medium to fine 1' 0" 

Sample 18. Channel sample of all of unit 5. 
4. Mixture of quartz sand, clay, and fine phosphorite. Yellow- 
brown to brown. 

Has appearance of weathered marl. No recognizable bedding fea- 
tures. Contains casts and molds of marine mollusks. Some 
silicified oyster shells. Occasional manatee ribs. 

Sand-size phosphorite is whitish, tan, brown, and orange. 

Quartz sand is fine to medium 1' 4" 

Sample 19. Channel sample of all of unit 4. 
3. Mixture of quartz sand, carbonate, clay, and sand-size phosphate 
particles. White with local white and yellow-brown mottling. 

No clear bedding features. Resembles a weathered marl. 

Phosphorite is white to tan to brown and orange. 

Quartz sand is medium to fine. 

Occurs as a thin lens 5" 

Sample 20. Channel sample of all of unit 3. 
2. Mixture of quartz sand, carbonate, clay, and sand-size phosphor- 
ite. Tan to yellow to yellow-brown. Gray where unweathered. 

Quartz sand is fine to coarse. 

Phosphorite is abundant and ranges in color from whitish to tan 
to brown and orange. Much brown and orange. 



50 Quarterly Journal of the Florida Academy of Sciences 

No clear bedding features in this unit I'll" 

Sample 21. Channel sample of upper 17 inches of unit 2. 

Bed Rock of the Hawthone Formation 

1. Pale yellow to yellow-brown limestone or dolomite. 

Quartz sand and brown to orange sand-size phosphorite dissem- 
inated through unit. 

Locally, there are thin stringers and fingers consisting almost 
entirely of quartz sand and fine brown to orange phosphorite. 

Quartz sand is fine to medium. 

At site of section only 8 inches of this rock is exposed. However, 
approximately 50 feet from the site of the measured section a 
drainage ditch has been cut to a depth of 4 feet into the sedi- 
ments at the bottom of the pit. In this ditch an additional 4 
feet of the pale yellow bed rock is exposed 8" 

Sample 22. Spot sample of pale yellow bed rock of unit 1. 



Total depth 31' 7" 

* Measured by E. C. Pirkle and W. H. Yoho, December, 1962. For 
analyses of samples see Tables 7-9. 

Achan Phosphate Mine 

Section 33, T. 30 S., R. 23 E., Polk County, Florida, 
approximately 4 miles southwest of Mulberry 

Thickness 
(in feet 
Unit Description and 

inches) 



Surface Sands 

10. Surface sands, loose. Very fine to medium, white at surface, 

grading downward into dark organic zone near base 3' 6" 

Sample 1. Spot sample of white sand collected 1 foot 5 inches 

below land surface. 
Sample 2. Spot sample taken at a depth of 2 feet 3 inches be- 
neath land surface in darker sands. 
Sample 3. Spot sample from dark organic zone. Collected at 
depth of 3 feet below land surface. 

Phosphatic Sediments of the Bone Valley District 

9. Clayey sand. "Sandrock" in places. Leached zone. Quartz sand 
is coarse to very fine. Whitish. Locally openings are present 
and suggest that material has been removed through weathering. 

Rare grains of whitish phosphorite 6' 10" 

Sample 4. Spot sample taken 1 foot 8 inches down from upper 
surface of unit. 



Pirkle, Yoho, Allen: Analyses of Sediments 51 

Sample 5. Spot sample collected 3 feet 4 inches above base of 
unit. 

Sample 6. Spot sample from lower 10 inches of unit. 

Sample 7. Channel sample from all of unit except bottom 10 
inches. 
8. Sand, clayey, to clayey sand. Quartz sand is medium to very fine. 
Whitish to light gray. 

Rare white phosphate particles 1' 7" 

Sample 8. Spot sample taken 7 inches below upper surface of 
unit. 

Sample 9. Spot sample from near base of unit. 
7. Sandy clay. Sand is medium to very fine. Clay is light gray to 
greenish-gray. 

A few soft white grains of phosphorite are present. 

Unit occurs as a thin lens that can be traced for several tens of 
feet along the mine face. 

Middle of lens is relatively hard and apparently has been silici- 
fied or phosphatized. Locally this hardened material is vesicular 
with soft white phosphate or phosphatic clay present in some 
partly developed openings 4" 

Sample 10. Spot sample of unit. 
6. Sand, clayey. Quartz sand is medium to very fine. White to 
light gray. 

A few white to gray phosphate particles. 

Present are stringers and small lenses containing a relatively high 
percentage of clay. 

Some workers would consider units 6 upward through unit 9 as 
the upper sand to clayey sand zone of the Bone Valley Forma- 
tion 1' 5" 

Sample 11. Channel sample of all of unit 6. 
5. Mixture of phosphate particles, quartz sand and clay. Whitish to 
gray. Quartz sand is fine to coarse. 

Pebble-size and granule-size phosphorite is very abundant. Sand- 
size phosphorite is common. Phosphate particles are white to 
gray to black. White colors predominate in upper part of unit; 
black colors are dominant lower in unit. 

Stringers and thin lenses of more massive greenish-gray clay are 
present throughout unit. 

This unit is separated from underlying unit 4 by a thin lens of 
massive gray to greenish-gray clay. This lens, approximately 2 
inches in thickness, was not sampled and therefore is not re- 
flected in any of the analyses. 

This unit is the uppermost one containing abundant phosphate 

particles 2' 3" 

Sample 12. Spot sample from near top of unit 5. 

Sample 13. Spot sample taken 1 foot 2 inches above base of 
unit 5. 



52 Quarterly Journal of the Florida Academy of Sciences 

Sample 14. Spot sample from near base of unit 5. 
4. Mixture of phosphate particles, quartz sand, and clay. Quartz 
sand is medium to very fine. Light blue-gray color where un- 
weathered, light tan to buff where weathered. 

Pebble-size and granule-size phosphorite is abundant. Sand-size 
phosphorite is common. In general the pebble and granule-size 
phosphorite is more abundant in upper part of unit. 

Phosphate particles are whitish, gray and black. The whitish 
particles are more abundant in upper part of unit. Black colors 
become more dominant lower in unit. 

Upper 5 inches of unit characterized by an increase in pebble- 
size phosphorite. A few of the pebbles are as much as 2 inches 
in longest dimension 2' 1' 

Sample 15. Spot sample from top 5 inches of unit. 

Sample 16. Spot sample from middle of unit. 

Sample 17. Spot sample from lower 9 inches of unit. 
3. Mixture of quartz sand, phosphorite and clay. Quartz sand is 
medium to very fine. Gray where unweathered, buff where 
weathered. 

Unit is different from overlying unit 4 and underlying unit 2 be- 
cause of its lack of abundant pebble-size and granule-size phos- 
phorite. This unit is characterized by its abundant sand-size 
phosphate particles. 

Phosphorite is mostly black. Some gray and whitish. Near top 

of unit whitish colors are slightly more common 8' 5" 

Sample 18. Spot sample from near top of unit. 

Sample 19. Spot sample taken 3 feet 6 inches below upper sur- 
face of unit. 

Sample 20. Spot sample from near base of unit. 
2. Mixture of phosphorite, quartz sand, and clay. Quartz sand is 
medium to very fine. Gray to light bluish-gray. Weathers to 
light tan or buff. 

Abundant pebble-size, granule-size and sand-size phosphate par- 
ticles. Phosphate pebbles are black, brown, white, and gray. 
Phosphate grains are gray, tan, brown, orange, white, and black. 

Near top of unit some phosphate pebbles are as much as 2 inches 
in longest dimension. 

A 3 inch thick zone of brown or drab colored material relatively 
high in clay is present at the top of this unit. 

Some workers would designate units 2 through 5 as the lower phos- 
phorite zone of the Bone Valley Formation 14' 7" 

Sample 21. Spot sample of drab material at top of unit. 

Sample 22. Spot sample collected near top of unit. 

Sample 23. Spot sample taken 3 feet below upper surface of unit. 

Sample 24. Spot sample taken 6 feet 6 inches down from top of 
unit. 

Sample 25. Spot sample collected 5 feet above base of unit. 



Pirkle, Yoho, Allen: Analyses of Sediments 53 

Sample 26. Spot sample selected from sediments near base of 
unit. 

Bed Kock of the Hawthorne Formation 

1. Pale yellow bed rock. Limestone, gray to bluish-gray where un- 

weathered. Weathers to pale yellow, light tan, and buff colors. 
Locally quartz sand, clay, and phosphate particles are important 

constituents of the limestone. Quartz sand is medium to very 

fine. 
Phosphate particles are brown to orange. 
In places this pale yellow limestone or dolomite is highly fossilif- 

erous (numerous impressions of marine mollusks). 

Only upper 4 inches of this limestone exposed 4" 

Sample 27. Spot sample of pale yellow bed rock. 



To bed rock and water 40' 4" 

* Measured by E. C. Pirkle and W. H. Yoho, November, 1962. For 
analyses of samples see Tables 10-12. 

Clermont Sand Mine of the Davenport Sand Company 

Section 27, T. 22 S., R. 26 E., Lake County, Florida, 
approximately 4 miles east-southeast of Clermont * 

Thickness 
(in feet 
Unit Description and 

inches) 

Surface Sands 

25. Surface sands, loose. Medium to fine, white, light tan, light brown 

to various shades of orange 3' 5" 

Citronelle Formation 

24. Clayey sand. Fine to coarse, reddish-brown. 
Bedding is essentially horizontal. 

Some stringers and small lenses contain very coarse quartz sand, 
quartz, or quartzite granules and discoid quartzite pebbles. The 
largest pebbles noted measured 1V2 inches in longest dimension. 
Occasional white kaolin balls disseminated through unit. 

Hardened or indurated through weathering 5' 6" 

23. Clayey sand. Very coarse to fine, orange-brown. 

Highly cross-bedded, scour type. Several different sets of cross- 
laminations. 
Hardened through weathering. 

Cross-bedding readings made in two different sets of cross-lamina- 
tions. 



54 Quarterly Journal of the Florida Academy of Sciences 

(First reading, greatest dip: 7 degrees to N. 55 E.) 

(Second reading, greatest dip: 11 degrees to S. 48 W.) 2'11" 

22. Clayey sand. Fine to very coarse, light brown to mottled white 
and brown. 

Small quartzite pebbles present near base of unit. 

Hardened through weathering. 

Units 22, 23 and 24 constitute the red and yellow zone 2' 1" 

21. Sand, clayey. Sand very coarse to very fine. White, locally mot- 
tled white, yellow, and orange. 

No clear sedimentary features. However, the sediments are prob- 
ably horizontally bedded and thinly laminated. 

Quartz or quartzite granules are more abundant near base of 
unit; clay content increases in upper few inches of the sedi- 
ments. 

Slightly hardened through weathering I'll" 

20. Sand, clayey. Sand very coarse to fine. White with orange bands 
Vs to x k inch thick spaced V2 to 3 inches apart. 

Highly cross-bedded, scour type. Two sets of cross-laminations 
dominant, a major upper set 1 foot 2 inches thick and a minor 
lower set 5 inches thick. In the lower set the sand is coarser, 
and quartz or quartzite granules are conspicuous. 

Units 20 and 21 are the major transitional units between over- 
lying red and yellow zone and underlying more dominantly 
white zone. 

Cross-bedding readings: 

(Major upper set, greatest dip: 13 degrees to N. 54 W.) 

(Minor lower set, greatest dip: 15 degrees to S. 10 W.) 1' 7" 

19. Sand. Medium to very coarse, white to dark brown. 

Horizontally bedded. 

Top 4 inches is white and contains a noticeable increase in clay. 

Much of the middle part of the unit is white, with occasional 
orange bands V* inch thick. This middle portion is low in clay, 
high in coarse sand, and is free-falling. 

Basal 9 inches is dark brown to maroon; the clay content is rela- 
tively high, and the sediments are lithified 2' 4" 

18. Clayey sand. Very fine, fine, and coarse, light pink to lavender. 

Although bedding features are not clearly visible, the sediments 

probably are horizontally bedded and thinly laminated I'll" 

17. Sand, clayey. Sand is very coarse, coarse, fine, and very fine. 
Materials are light pink to lavender. 

Numerous stringers and small lenses of coarse sediments are in- 
tercalated with finer sands that contain a relatively high clay 
content. Stringers and lenses of coarse sediments are more nu- 

numerous near base of unit 2' 2" 

16. Clayey sand. Very fine, fine, and coarse, light lavender to pink 
to mottled lavender, pink and white. 



Pirkle, Yoho, Allen: Analyses of Sediments 55 

Thinly laminated and horizontally bedded. Alternate laminations 
of coarse sediments and fine sediments. The fine sediments 
contain more clay. 
Numerous small kaolin balls about the size of a pin head dissem- 
inated through unit. Such small kaolin balls probably result 
from the fragmentation of thin clay seams. 

Relatively high in white mica 2' 3" 

15. Sand. Very coarse to coarse. White with tints of crimson and 
orange. 
Noticeable quartz or quartzite granules and a few quartzite peb- 
bles with longest dimensions as much as Yz inch. 
Occurs as a small lens of coarse sediments separating overlying and 
underlying finer and more clayey units. This lens ranges in 
thickness from about 1 inch to a foot and can be traced for sev- 
eral tens of feet along the pit face 7" 

14. Clayey sand. Very fine to coarse, mottled lavender and white. 
Bedding features obscured. 

Occasional tiny kaolin balls disseminated through unit 1' 3" 

13. Sand, clayey. Sand fine to medium, white with a light pink cast. 
Lower part of unit shows distinct cross-bedding. 

Cross-bedding reading (greatest dip: 8 degrees to N. 20 W.) 7" 

12. Sand. Very coarse to fine, white. 

Present as a thin lens 4" 

11. Sand, clayey. Sand very fine to coarse, white with a pink (almost 
salmon) tint. 

A small lens 3" 

10. Sand. Very coarse to fine, white. 

Horizontally bedded and thinly laminated. 

Bedding is very distinct for a coarse unit 1' 1" 

9. Sand, clayey. Sand very coarse to fine, white toward top, lav- 
ender toward base. 
Faint cross-bedding but could not measure with confidence the 

direction of greatest dip of cross-laminations V 0" 

8. Clayey sand. Very fine, fine and coarse, white. 

No clear bedding features but materials are probably horizontally 
bedded and thinly laminated. 

Mica content relatively high 11" 

7. Sand. Coarse to fine, white with slight pink and lavender cast. 

Occurs as a small lens T 7" 

6. Clayey sand. Fine to very fine, white. 

A few stringers containing coarse to very coarse sand are present 
in the unit. One small lens of coarse sediments is present near 
middle of unit. This lens is about Yz inch thick 5" 

5. Sand. Very coarse to fine, white with some lavender and pinkish- 
lavender banding going with cross-laminations. Bands are Ys 
to 1 inch thick. 



56 Quarterly Journal of the Florida Academy of Sciences 

Unit highly cross-bedded, scour type. Cross-laminations cut off 
sharply at top of unit. Contact with underlying unit distinct. 

Sediments are thinly laminated. Most laminae Vs to V2 inch thick. 
The coarse laminations are more pronounced. 

Numerous small white kaolin masses and balls, size of pin head 
to Va inch in diameter disseminated through unit. Thin clay 
seams in various stages of fragmentation. 

Cross-bedding readings: 

(First reading, about 8 inches down from upper surface of unit, 
greatest dip: 15 degrees to N. 20 W.) 

(Second reading, taken directly beneath first reading and just over 
base of unit, greatest dip: 17 degrees to N. 15 W.) 

(Third reading, at same stratigraphic posititon as first reading but 
2 feet to east in a different cross-bedded set, greatest dip: 14 
degrees to N. 12 E.) 

(Fourth reading, taken directly beneath third reading and just over 

base of unit, greatest dip: 10 degrees to N. 16 E.) 1' 5" 

4. Clayey sand. Fine to medium, mottled white and pinkish-laven- 
der. 

Horizontally bedded. 

Numerous thin clay seams (mainly horizontal) only a fraction of 
an inch to slightly more than an inch in length and from 

% to Va inch thick 2' 6" 

3. Sand, clayey. Sand fine to medium, white to slightly pinkish- 
lavender cast. 

No clear bedding features 5' 0" 

2. Sand, clayey. Sand fine to coarse, whitish 9" 

1. Covered interval to lake level 5' 1" 



To lake level 47'10" 

* Measured by E. C. Pirkle, Fred Pirkle, and W. H. Yoho, late 1962 and 
early 1963. Analyses of channel samples of each unit are given in Tables 
13-15. 

Literature Cited 1 

Altschuler, Z. S., E. B. Jaffe, and Frank Cuttita. 1956. The aluminum 
phosphate zone of the Bone Valley formation, Florida, and its uranium 
deposits. U. S. Geol. Survey Prof. Paper 300, pp. 495-504. 

Altschuler, Z. S., and E. J. Young. 1960. Residual origin of the "Pleisto- 
cene" sand mantle in central Florida uplands and its bearing on marine 



1 This paper was accepted for publication on November 6, 1964. A 
paper on "Stratigraphy and sedimentation of phosphorite in the central Flor- 
ida phosphate district" was read by Stanley R. Riggs and Donald H. Freas 
at the annual meeting of the American Institute of Mining, Metallurgical, and 
Petroleum Engineers, Chicago, February 14-18, 1965, too late for considera- 
tion here. 



Pirkle, Yoho, Allen: Analyses of Sediments 57 

terraces and Cenozoic uplift. U. S. Geol. Survey Prof. Paper 400-B, 
pp. B 202-B 207. 

Bishop, Ernest W. 1956. Geology and ground water resources of High- 
lands County, Florida. Florida State Geol. Surv., Rept. Invest., no. 
15, pp. 1-115. 

Brodkorb, Pierce. 1955. The avifauna of the Bone Valley formation. Flor- 
ida State Geol. Surv., Rept. Invest., no. 14, pp. 1-57. 

. 1963. Miocene birds from the Hawthorne Formation. Quart. Jour. 

Florida Acad. Sci., vol. 26, pp. 159-167. 

Carr, W. J., and D. C. Alverson. 1959. Stratigraphy of middle Tertiary 
rocks in part of west-central Florida. U. S. Geol. Surv., Bull. 1092, 
pp. 1-111. 

Cathcart, J. B. 1950. Notes on the land-pebble phosphate deposits of 
Florida, in Symposimn on mineral resources of the southeastern United 
States. University of Tennessee Press, pp. 132-151. 

. 1963a. Economic geology of the Keysville Quadrangle Florida. 

U. S. Geol. Surv. Bull. 1128, pp. 1-82. 

. 1963b. Economic geology of the Chicora Quadrangle Florida. 

U. S. Geol. Surv. Bull. 1162-A, pp. A 1- A 66. 

. 1963c. Economic geology of the Plant City Quadrangle Florida. 

U. S. Geol. Surv. Bull. 1142-D, pp. D 1- D 56. 

Cooke, C. Wythe. 1945. Geology of Florida. Florida State Geol. Surv., 
Geol. Bull., no. 29, pp. 1-339. 

Espenshade, G. H., and C. W. Spencer. 1963. Geology of phosphate de- 
posits of northern peninsular Florida. U. S. Geol. Surv., Bull. 1118, 
pp. 1-115. 

Hunter, Frank R. 1949. Occurrence of heavy minerals in the pebble 
phosphate deposits of Florida. A.I.M.E. Trans., vol. 181, pp. 413-416. 

Isphording, W. C. 1963. A study of the heavy minerals from the Haw- 
thorn Formation and overlying Pleistocene sands exposed at the Devil's 
Mill Hopper, Alachua County, Florida. University of Florida, un- 
published thesis, pp. 1-31. 

Ketner, K. B., and L. J. McGreevy. 1959. Stratigraphy of the area be- 
tween Hernando and Hardee counties, Florida. U. S. Geol. Surv. Bull. 
1074-C, pp. 49-124. 

Klein, Howard, M. C. Schroeder, and W. F. Lichtler. 1964. Geology 
and ground-water resources of Glades and Hendry counties, Florida. 
Florida State Geol. Surv., Rept. Invest., no. 37, pp. 1-101. 



58 Quarterly Journal of the Florida Academy of Sciences 

Martens, James H. C. 1935. Beach sands between Charleston, South Caro- 
lina, and Miami, Florida. Geol. Soc. America Bull., vol. 46, pp. 
1563-1596. 

McClellan, Guerry H. 1962. Identification of clay minerals from the 
Hawthorn Fonnation, Devil's Mill Hopper, Alachua County, Florida. 
University of Florida, unpublished thesis, pp. 1-38. 

PmKLE, E. C. 1956. The Hawthorne and Alachua formations of Alachua 
County, Florida. Quart. Jour. Florida Acad. Sci., vol. 19, pp. 197- 
240. 

. 1958. Lithologic features of Miocene sediments exposed in the 

Devil's Mill Hopper, Florida. Quart. Jour. Florida Acad. Sci., vol. 
21, pp. 149-161. 

Peeikle, E. C, and W. H. Yoho. 1961. Folding and warping resulting from 
solution with associated joints and organic zones in clayey sands at 
Edgar, Florida. Quart. Jour. Florida Acad. Sci., vol. 24, pp. 247-266. 

Pirkle, E. C, W. H. Yoho, A. T. Allen, and A. C. Edgar. 1963. Citro- 
nelle sediments of peninsular Florida. Quart. Jour. Florida Acad. Sci., 
vol. 26, pp. 105-149. 

Pirkle, E. C, W. H. Yoho, and A. T. Allen. 1964. Origin of the silica 

sand deposits of the Lake Wales Ridge area of Florida. Econ. Geol., 

vol. 59, pp. 1107-1139. 
Puri, Harbans S., and Robert O. Vernon. 1959. Summary of the geology 

of Florida and a guidebook to the classic exposures. Florida State 

Geol. Surv., Special Pub. 5, pp. 1-255. 

Reves, William D. 1960. An X-ray study of two Florida land pebble 
phosphate samples, in 9th Field Trip Guide Book of Southeastern 
Geological Society, Tallahassee, pp. 50-63. 

Sellards, E. H. 1909. Mineral industries. Florida State Geol. Surv., Sec- 
ond Ann. Rept., pp. 235-291. 

. 1910. A preliminary paper on the Florida phosphate deposits. 

Florida State Geol. Surv., Third Ann. Rept., pp. 17-41. 

Simpson, G. G. 1929. The extinct land mammals of Florida. Florida State 
Geol. Surv., Twentieth Ann. Rept., pp. 229-279. 

Thoenen, John R., and John D. Warne. 1949. Titanium minerals in cen- 
tral and northeastern Florida. U. S. Bur. Mines Rept. Inv. 4515, pp. 
1-62. 

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Florida State Geol. Survey Bull., 33, pp. 1-256. 

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Georgia. 

Quart. Jour. Florida Acad. Sci. 28(1) 1965 



A NEW CIRRIPED FROM THE EOCENE OF GEORGIA 

Arnold Ross 

Late in 1964 Dr. Katherine Van Winkle Palmer, of the Paleon- 
tological Research Institution, forwarded to the author a large 
number of disarticulated, and a few articulated, balanomorph 
barnacles from the Eocene of Georgia. Eocene Balanidae from 
the Atlantic and Gulf Coastal Plains are virtually unknown. 

In the appendix to his "Synopsis of the organic remains of the 
Cretaceous group" Samuel Morton (1834) cited Balanus ostrearum 
as a member of the southeastern United States fauna. Both Dar- 
win (1854b), and Kolosvary (1955, 1961) attributed this species to 
Conrad as did Morton. Conrad may have found the specimen, 
assigned a name to it, and allowed Morton to publish it; Conrad 
never published the name. Because this species was neither de- 
scribed nor figured the author must therefore consider it to be a 
nomen nudum. 

Morton (1834, p. 72), earlier in the same paper, reported a 
species "Found by Mr. Conrad in the calcareous strata of South 
Carolina," which he named Balanus peregrinus. Darwin (1854b, 
p. 492), in referring to B. peregrinus, stated, "... this Eocene 
species apparently resembles the B. unguiformis of Sowerby . . . " 
This appears unlikely since B. peregrinus is a well-ribbed form that 
reaches a " . . . diameter from half an inch to one inch and a half" 
(Morton, 1834, p. 72). Although this species was originally de- 
scribed and figured by Morton, and subsequently by Conrad 
(1846a), its true nature cannot be determined at this time because 
the diagnostic characters cited are presently of no taxonomic value. 
Balanus peregrinus is here considered as a species inquierenda. 

Balanus humilis was described by Conrad (1846b) from a lime- 
stone outcropping in the Tampa Bay region ("Hillsborough Falls") 
of Florida. This species was believed to be of early Eocene age, 
although at present the Tampa Limestone, from which this species 
presumably was collected, is assigned an early Miocene age. 

In the obscure half-page paper by Holmes (1859) two balanids 
were reported from the "Eocene Marl of Ashley River," Balanus 
digitatus and Balanus calceolus. Balanus calceolus Ellis is a Eu- 
ropean conopean that ranges from Helvetian to Recent. The 
short, one-paragraph descriptions of the species unfortunately do 
not lend themselves to modern studies. Holmes' failure to obtain 



60 Quarterly Journal of the Florida Academy of Sciences 

opercular valves further obscures the true nature of both forms. 
His illustrations add nothing to terse, but unimportant morpho- 
logical details. Furthermore, the stratigraphic occurrence of these 
species must be redetermined in the light of subsequent refine- 
ments in the stratigraphy of South Carolina. Therefore, it appears 
best to consider these two forms as species inquirendae. 

Otto Meyer (1886, 1887) reported "Balanus antiquus" originally 
described as a new species of Crucibulum (Gastropoda), from the 
Claiborne Formation, middle Eocene, of Alabama. Although Pils- 
bry (1930) contended that it was "not recognizably defined," he 
did note that the opercular valves of the sole specimen were pres- 
ent. Zullo (letter dated November 29, 1964) found that several 
specimens in the University of California Museum of Paleontology 
from the Claiborne and Jackson groups in Alabama may be refer- 
able to "antiquus." 

Withers (1953) reported the presence of Balanus sp. aff. B. un- 
guiformis Sowerby from sediments of the upper Eocene Jackson 
group of Mississippi. The stratigraphic range of this species is 
Auversian (Zullo, 1960b) through Rupelian (Withers, 1953), al- 
though Davadie (1963) recorded it from sediments as late as Hel- 
vetian. Zullo (1960a) reported the occurrence of a species con- 
specific with, or related to, B. unguiformis from the Claiborne and 
Jackson groups of Alabama. He has also (1960b) noted the oc- 
currence of an undescribed hesperibalanid on the Pacific Coast of 
the United States. 

Several Eocene Balanidae have been described from European 
sediments. The hesperibalanids Balanus unguiformis and B. eris- 
ma, described by James De Carle Sowerby in 1846, were probably 
the first Eocene species reported from Europe. Darwin (1854a) 
considered B. erisma to be a variety (= subspecies) of B. ungui- 
formis, and further considered (Darwin, 1854b) B. perplexus Nyst, 
1853, from the Eocene of Belgium, to be conspecific with B. un- 
guiformis. Kolosvary (1947) reported two species from Hungary, 
B. hantkeni from the middle Eocene, and B. phineus from the late 
Eocene (1958). Both species were, unfortunately, described solely 
on the basis of shell morphology. Recently, Kolosvary (1961) de- 
scribed another balanid, B. vialovi, from the late Eocene of the 
U.S.S.R. In addition, members of the B. concavus and B. tin- 
tinnabulum complexes have been reported from Eocene sediments 
(Davadie, 1963), but these records must be substantiated. 



Ross: New Eocene Barnacle 61 

Balanus sublaevis was briefly described and figured by Sowerby 
in a geological memoir by Grant (1840) and has not since been 
redescribed. Recently, Kolosvary (1961) cited this Asian species 
as an Eocene faunal element, even though it was originally re- 
ported from "Tertiary" sediments. It is the writers belief that 
Kolosvary's assignment is unjustifiable. 

Institutional abbreviations used in the present study are as 
follows: Paleontological Research Institution, P.R.I. ; Florida State 
Museum, F.S.M. 

Type Locality 

The sediments from which the barnacles were collected outcrop 
approximately 22 miles southeast of Augusta on the Savannah 
River, at Shell Bluff Landing, Burke County, Georgia (P.R.I, field 
station 1894). Maps of this area may be found in Herrick's studies 
on the Shell Bluff Foraminifera (1960, 1964). 

A generalized lithologic section at this locality was presented 
by Cooke (1943). The barnacles were collected from Cooke's 
unit 6, which is referred to the Barnwell Formation of Eocene 
(Jacksonian) age. Herrick (1964) confirmed Cooke's age determin- 
ation, and correlated this horizon with "... the type Moodys 
Branch Formation of Mississippi." 

The specimens herein described were collected by the late 
G. D. Harris, J. Hauck, and Katherine V. W. Palmer in October, 
1946. 

Order THORACICA Darwin, 1854 

Family Balanidae Gray, 1825 

Subfamily Balaninae (Gray), 1825 

Genus Kathpalmeria, new genus 

Definition. Sessile barnacles having six compartments with 
solid walls and solid, wholly calcareous, basis. The rostrum, lat- 
erals, and carinolaterals lack (?) or possess diminutive radii, the 
sutural edges of which are dentate. The wall plates are mod- 
erately folded, the reentrants forming buttresses on the inner shell 
surface. Internally, the buttresses and intervening spaces are 
strong to slightly ribbed. The scutum is sulcate externally, and 
bears a strongly developed articular ridge, but no adductor ridge. 



62 Quarterly Journal of the Florida Academy of Sciences 

Etymology. This new genus is named in honor of Dr. Kath- 
erine Van Winkle Palmer in recognition of her outstanding con- 
tributions to the Eocene paleontology of the southeastern United 
States. 

Type species. Kathpalmeria georgiana, new species. 





Fig. 1. Kathpalmeria georgiana, new genus, new species. Drawings of 
external and internal views of holotype scutum, P.R.I, no. 6075b. Actual 
height 4.3 mm. Drawings prepared by Miss Brenda Baer. 

Remarks. This new genus is proposed to include, in addition 
to the new species described below, Balanus hantkeni Kolosvary 
(1947) from the middle Eocene of Hungary. In many respects both 
of these species are closely related to Armatobalanus and Hesperi- 
balanus, subgenera of the genus Balanus. Strongly, moderately, or 
slightly ribbed forms appear in both of these groups, but the rib- 
bing is not reflected internally as in Kathpalmeria. All of the pres- 
ently known hesperibalanids possess well-developed radii which 
bridge completely the gap between the parietes; with one excep- 
tion the same feature appears in all of the armatobalanids. In 
Recent hesperibalanids there is a well to poorly developed ad- 
ductor ridge; fossil forms also show various degrees of develop- 
ment of this structure. On the other hand, certain of the armato- 
balanids, namely, B. nefrens and quadrivittatus, do not have an 
adductor ridge. The solid nature of the shell and basis, unusual 
folded walls, obsolete or diminutive radii, and the scutum with 
no adductor ridge should serve to distinguish this new genus from 
all of the presently known genera of the subfamily Balaninae. 



Ross- New Eocene Barnacle 



63 




Fig. 2. Kathpalmeria georgiana, new genus, new species, a, b, external 
and internal views of right lateral, paratype, P.R.I, no. 6076, actual height 
10 mm. c, external view of partially disarticulated shell, paratype, F.S.M. 
no. 4000, actual height 10.1 mm. d, internal view of right lateral, paratype, 
P.R.I, no. 6077, actual height 14.3 mm. e, internal view of rostrum, paratype, 
P.R.I, no. 6078, actual height 10.5 mm. /, internal view of right lateral, 
paratype, P.R.I, no. 6079, actual height 13.3 mm. 



Kathpalmeria georgiana, new species 

Description. The shell is low conic and of a moderate size. 
The mean height of six articulated specimens is 7.6 mm. The walls 
are moderate to strongly folded, the reentrants forming buttresses 
on the inner surface. Growth ridges on the external surface ap- 
pear best developed on the reentrants. Viewed from the basal 
edge the parietal plates appear sinuous. The peritreme is but 
slightly toothed. The orifice is pentagonal in shape, being widest 
at the rostral end. The exterior of the parietes are ornamented 
with strong, moderately narrow to broad, non-digitating ribs; the 



64 Quarterly Journal of the Florida Academy of Sciences 

ribs extend from the base to the apex. The ribs are separated by 
approximately their own width or greater. On some plates there 
are intercalated secondary ribs. Under magnification alternating 
dark and light vertical bands of shell material can be seen. Upon 
weathering of the shell there is evidently a differential degree of 
solubility of these bands since the light bands are eroded, and the 
dark ones present the appearance of uniform secondary riblets. 
The radii are extremely narrow to obsolete, do not extend the 
complete height of the valve, and the sutural edges are septate. 
The septa appear as simple, square pegs; there are no denticles 
on either the top or bottom edges of the septa. The alae 
have horizontal summits, and the sutural edges of the alae are 
also septate. Internally, the inner shell surface is marked by sub- 
sidiary ribs on the inner manifestations of the ribs and reentrants. 
The ribs may extend to the hollow behind the sheath. The sheath 
surface is strongly scored, and the growth ridges are apically di- 
rected. The basal margin of the sheath is either free or attached. 
Where free, the hollow behind it is narrow, and moderately deep. 

The surface of the flat basis is smooth. At the periphery den- 
ticles are developed which interlock with those developed on the 
basal edges of the wall plates. The wall plates and basis are ap- 
parently strongly articulated since most of the disarticulated com- 
partments still retain peripheral portions of the basis. 

The scutum is broadly triangular, and strongly concave be- 
tween the basal margin and apex. The basal margin is slightly 
longer than the tergal margin. The occludent margin is not pecti- 
nate. The apex of the valve is broadly rounded. It cannot be de- 
termined at this time whether this is due to secondary erosion. A 
somewhat prominent sulcus extends from just left of the center 
of the apex and extends down the occludent side to the basal mar- 
gin dividing the valve unequally. The exterior ornamentation con- 
sists of strong, spaced, growth ridges. It appears that the growth 
ridges may have been crossed by numerous, close-spaced longitudi- 
nal striae. The articular ridge is high, slightly reflexed, evenly 
arched, approximately two-thirds the length of the tergal margin, 
and distally terminates abruptly. The articular furrow is extremely 
broad, and moderately deep. There is no adductor ridge. The ad- 
ductor muscle pit is large, somewhat above center, deep, and well 
defined apically, but open below. The inner face of the articular 
ridge, and possibly the apical portion of the valve proper, bear 



Ross: New Eocene Barnacle 65 

prominent, rectilinear, parallel ridges. The pit for the lateral de- 
pressor muscle is neatly triangular, extremely deep, and extends 
apically about one-fourth the height of the valve. The mediad 
limits of the rostral depressor muscle pit are poorly defined, but 
the pit is deep and extends a short distance above the basal 
margin. 

Disposition of Types. The holotype right scutum (fig. 1) and 
associated shell (not figured) are deposited in the collections of 
the Paleontological Research Institution, catalog numbers 6075b 
and 6075a, respectively. The paratypic material, with one ex- 
ception as noted below, is deposited with the above institution, 
catalog numbers 6076 (right lateral, fig. 2a, b), 6077 (right lateral, 
fig. 2d), 6078 (rostrum, fig. 2e), 6079 (right lateral, fig. 2f), and 6080 
(not figured). One partially disarticulated specimen (fig. 2c) has 
been placed in the Florida State Museum collections, catalogue 
number 4000. 

Measurements of Holotype. Height of shell 8.1; carinorostral 
diameter of shell 11.5; carinorostral diameter of orifice 5.1; height 
of right scutum 4.3 mm. 

Acknowledgments 

The author is grateful to Dr. Katherine Van Winkle Palmer for 
permission to study the barnacles described herein. Special thanks 
are due Dr. Victor A. Zullo, Marine Biological Laboratory, Woods 
Hole, for his invaluable criticisms and comments on the manu- 
script. Dr. Steve M. Herrick of the Atlanta Ground Water Branch, 
U. S. Geological Survey, kindly furnished the author with copies 
of his papers on the Shell Bluff Foraminifera. The assistance of 
Mr. Ray Jones of the Reference and Bibliography section, Uni- 
versity of Florida Libraries, in obtaining rare publications during 
the course of the present and many earlier studies is gratefully 
acknowledged. 

Literature Cited 

Conrad, Timothy A. 1846a. Observations on the Eocene formation of the 
United States, with descriptions of species of shells, &c. occurring in 
it. Amer. Jour. Sci. Arts, ser. 2, vol. 1 (for 1845), no. 2, pp. 209-221. 

. 1846b. Descriptions of new species of organic remains from the 

upper Eocene limestone of Tampa Bay. Ibid., ser. 2, vol. 2, no. 6, 
pp. 399-400. 



66 Quarterly Journal of the Florida Academy of Sciences 

Cooke, C. Wythe. 1943. Geology of the coastal plain of Georgia. Bull. 
U. S. Geol. Surv., no. 941, pp. 1-121. 

Darwin, Charles R. 1854a. A monograph on the fossil Balanidae and 
Verrucidae of Great Britain. London, Paleontographical Society, 44 
pp., 2 pis. 

. 1854b. A monograph on the sub-class Cirripedia, with figures of 

all the species. The Balanidae, (or sessile cirripeds); the Verrucidae, 
etc., etc., etc. London, Ray Society, 684 pp., 30 pis. 

Davadie, Claude. 1963. Systematique et structure des Balanes fossiles 
d'Europe et d'Afrique. Editions Centre National Recherche Scien- 
tifique, Paris, 146 pp., 57 text-figs., 55 pis. 

Grant, C. W. 1840. Memoir to illustrate a geological map of Cutch. Trans. 
Geol. Soc. London, ser. 2, vol. 5, pt. 1, pp. 289-329, pis. 20-26. Ap- 
pendix entitled: Systematic list of organic remains, the plants deter- 
mined by Mr. John Morris, and the remainder by Mr. James De Carl 
[sic] Sowerby, A.L.S. (pp. 327-329). 

Herrick, Steve M. 1960. Some small Foraminifera from Shell Bluff, Geor- 
gia. Bull. Amer. Paleon., vol. 41, no. 187, pp. 117-130, pis. 14-16, 
1 text-fig. 

. 1964. Upper Eocene smaller Foraminifera from Shell Bluff and 

Griffin Landings, Burke County, Georgia. U. S. Geol. Surv. Prof. 
Paper 501-C, pp. C64-C65, 1 text-fig. 

Holmes, Francis S. 1859. Descriptions of new fossil Balani, from the 
Eocene Marl of Ashley River, S. C. Proc. Elliott Soc. Nat. Hist, vol. 
1, p. 21, pi. 1, figs. 7-12. 

Kolosvary, G. 1947. Eine neue Balanidae aus dem ungarischen Eozan. 
Ann. Hist.-Nat. Mus. Nat. Hungarici, vol. 40, no. 8 3 pp. 305-307, text- 
fig. 1. 

. 1955. tiber stratigraphischer Rolle der fossilen Balaniden. Acta 

Biol. Szeged, new ser., vol. 1, pts. 1-4, pp. 183-188. 

. 1958. Phylogenetische Beitrage zur Gattung Balanus. Acta Zool., 

Acad. Sci. Hungaricae, vol. 2, pts. 1-3, pp. 187-191, text-fig. 1, pi. 1. 

. 1961. Further fossile balanids from the USSR. Acta Biol., new 

ser., vol. 7, pts. 3-4, pp. 149-154, text-figs. 1-3. 

Meyer, Otto. 1886. Contributions to the Eocene paleontology of Alabama 
and Mississippi. Bull. Alabama Geol. Surv., vol. 1, no. 2, pp. 63-85, 
pis. 1-3. 

. 1887. On invertebrates from the Eocene of Mississippi and Ala- 
bama. Proc. Acad. Nat. Sci. Philadelphia, vol. 39, pp. 51-56, pi. 3. 



Ross: New Eocene Barnacle 67 

Morton, Samuel G. 1843. Synopsis of the organic remains of the Creta- 
ceous group of the United States. To which is added an appendix, 
containing a tabular view of the Tertiary fossils hitherto discovered in 
North America. Philadelphia, Key and Biddle, 88 pp., 19 pis. Ap- 
pendix entitled: Catalogue of the fossil shells of the Tertiary forma- 
tions of the United States, embracing all the species hitherto pub- 
lished, 8 pp. 

Pilsbry, Henry A. 1930. Cirripedia (Balanus) from the Miocene of New 
Jersey. Proc. Acad. Nat. Sci. Philadelphia, vol. 82, pp. 429-433, text- 
figs. 1-2, pis. 36-37. 

Withers, Thomas H. 1953. Catalogue of fossil Cirripedia in the Depart- 
ment of Geology. Vol. 3. Tertiary. London, British Museum (Nat- 
ural History), 396 pp., 105 figs., 64 pis. 

Zullo, Victor A. 1960a. Cenozoic Balanomorpha of the Pacific Coast of 
North America. Unpubl. Univ. California (Berkeley) master's thesis, 
147 pp., 9 pis. 

. 1960b. Eocene species of the genus Balanus (Cirripedia). Bull. 

Geol. Soc. Amer., vol. 71, no. 12, pt. 2, p. 2084. 

Department of Geology, University of Florida, Gainesville, 
Florida. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



EARLY MIOCENE ANURANS FROM FLORIDA 
J. Alan Holm an 

The early Miocene beds of the Thomas Farm of Gilchrist 
County, Florida, have produced the most diverse anuran fauna 
known from the Tertiary of North America. Indeed, all six anu- 
ran families indigenous to Florida today are represented in these 
Miocene sediments. Previous references to Thomas Farm frogs 
and toads have been summarized by Holman (1961). A compre- 
hensive list of the Thomas Farm vertebrates with references is 
found in Olsen (1962). Since Olsen's paper, Auffenberg (1963) has 
discussed snake material, and Estes (1963) has reported on sala- 
mander and lizard fossils. Several of these authors have discussed 
the stratigraphy of the deposit. 

Large numbers of additional anuran bones have recently ac- 
cumulated through the intensive collecting of workers affiliated 
with the Florida Geological Survey and the University of Florida. 
These new bones have been submitted to me for identification, and 
the present report deals with the leptodactylid, ranid, and brevi- 
cipitid frogs of the deposit. A second report on the bufonid, hylid, 
and pelobatid frogs is anticipated as soon as enough recent skele- 
tons of these groups become available. 

Materials and Methods 

I have previously found the ilium to be a reliable element in 
the identification of fossil anuran remains (Holman, 1959, 1961, 
1962a, 1962b, 1963a, 1963b). Other workers have also found the 
ilium to be useful (see Auffenberg, 1956, 1957, 1958; Lynch, 1964; 
Tihen, 1962; and especially Chantell, 1964). I now regard the 
ilium as the best single element upon which to base the identifi- 
cation and description of fossil frogs, and I suggest that this ele- 
ment be designated as the type specimen when descriptions based 
on disarticulated anuran material are made. The sacrum, which 
Taylor (1942) favors, is quite subject to individual variation. In 
fact, it may show individual variation in characters that are con- 
sidered to be definitive at the subordinal level (Holman, 1963c). 
Nevertheless, the sacrum is a good postcranial element for the sub- 
stantiation of ilial identifications. Other postcranial elements are 
untrustworthy, and unfortunately, fossil anuran cranial elements 



Holman: Miocene Frogs 69 

are usually in the form of tiny fragments of disarticulated skull 
bones. 

Anuran skeletons at Illinois State University and material bor- 
rowed or received as gifts from individuals in the acknowledg- 
ments section have been utilized as comparative material. The 
following abbreviations are used: M.C.Z. — Museum of Compara- 
tive Zoology; U.F. — University of Florida; F.G.S. — Florida Geo- 
logical Survey. All measurements are in millimeters. 

Acknowledgments 

I wish to thank Bryan Patterson, Stanley J. Olsen, and Clayton 
Ray for the privilege of studying the fossil material in their care. 
Several persons have generously loaned or given comparative ma- 
terial used in this study. These people include: W. Auffenberg 
and P. Brodkorb, Gainesville, Florida; J. D. Lynch and H. M. 
Smith, Urbana, Illinois; W. F. Blair, Austin, Texas; R. S. Sim- 
mons, Baltimore, Maryland; G. B. Rabb, Brookfield, Illinois; and 
J. A. Peters and W. J. Riemer, Washington, D. C. Donna Rae 
Holman made the drawings. 

Family Leptodactylidae 

Heretofore, the only pre-Pleistocene record of the Leptodactyli- 
dae from North America is that of Estes (1964), who assigned a 
right squamosal to "Family incertae sedis, near Leptodactylidae?". 
Eleutherodactylus augusti and Syrrhophis marnocki have been re- 
ported from the Pleistocene of Texas (Mecham, 1959; Tihen, 1960; 
Lynch, 1964). 

Genus Leptodactylus Fitzinger 

In the study of leptodactylid fossils the following recent skel- 
etons were examined: Ceratophrys cornuta 1, C. varia 1, Eleuthero- 
dactylus augusti 1, E. dunni 5, E. podociferus 2, E. ricordi 2, Eu- 
pemphix pustulosa 4, Leptodactylus insularum 3, L. melanonotus 
5, Miobatrachus gouldi 1, Tomodactylus nitidus 2, and Syrrhophus 
marnocki 1. 

In concordance with the great adaptive radiation of the Lep- 
todactylidae their ilia show many striking intergeneric differences. 
The ilial crest is absent in Ceratophrys, Eupemphix, and Mioba- 
trachus. The other genera show an interesting convergence with 
some genera of the Ranidae in that an ilial crest is present. In 



70 Quarterly Journal of the Florida Academy of Sciences 

Eleutherodactylus dunni the crest is high throughout, but it is 
extremely thin and has a membranous appearance. In E. augusti, 
E. podociferus, E. ricordi, and Leptodactylus the crest is mod- 
erately high and is thin with the exception of E. augusti, which 
has a somewhat thicker crest. In Syrrhophus and Tomodactylus 
the crest is obsolete. There is great variability in the prominence 
for the origin of the vastus externus head of the M. triceps femoris 
(for the sake of brevity this structure will hereafter be referred to 
as the vastus prominence). In Ceratophrys it is a much produced 
dorsal spike, whereas in Miobatrachus it is obsolete. In Eleuthero- 
dactylus, Eupemphix, Syrrhophus, and Tomodactylus it is small 
to medium in size, ovaloid to round in shape, and moderately to 
strongly swollen. In Leptodactylus it is larger and is usually 
strongly beveled. 

Thus, the ilia of Leptodactylus may be distinguished from other 
Leptodactylidae at hand by the following combination of char- 
acters: ilium with crest present, moderately high, thin; vastus 
prominence large, usually strongly beveled. 

Figure 1 shows the distribution of muscular origins on an ilium 
of Leptodactylus melanonotus. The muscles of two Leptodactylus 
insularum specimens were also studied. Whenever possible I have 
tried to use terminology consistent with ilial musculature. Five 
ilia from the Thomas Farm represent a new species of Leptodac- 
tylus, the first fossil leptodactylid described from North America. 

Leptodactylus abavus sp. nov. 

Holotype. Right ilium, U.F. 10201 (Fig. 1). From Hawthorne 
formation, lower Miocene, Arikareean; Thomas Farm, Gilchrist 
County, Florida. Collected by Clayton Ray. 

Paratypes. Right ilium, F.G.S. V-6067; two right and one left 
ilia, U.F. 10202, from the same locality as the holotype. 

Referred elements. Two sacra, F.G.S. V-6068 and M.C.Z. 3406. 

Diagnosis. A small Miocene Leptodactylus showing similari- 
ties to recent L. melanonotus, but differing in having; posterodor- 
sal border of ilium sloping gently into dorsal acetabular expansion 
(L. melanonotus with this border sloping more abruptly into dorsal 
acetabular expansion); vastus prominence more produced and ex- 
tensive, with valley between it and dorsal acetabular border deep- 
er; angle between ventral acetabular expansion and shaft greater 



Holman: Miocene Frogs 



71 



than 90 degrees (L. melanonotus with angle less than 90 degrees). 
Measurements: greatest height through acetabulum 2.3; greatest 
height through ilial crest 2.2; greatest length of vastus prominence 
2.4 mm. 









Fig. 1. Top, holotype right ilium of Leptodactylus abavus sp. nov., U.F. 
10201 (line represents 2 mm.). Bottom, ilium drawn from Leptodactylus 
melanonotus to illustrate origin of selected muscles: A, acetabular fossa; B, 
tendinous origin of M. biceps femoris; C, fleshy origin of vastus externus head 
of M. triceps femoris; D, fleshy origin of M. gluteus; E, tendinous origin of 
M. sartorius; F, tendinous origin of vastus internus head of M. triceps femoris. 



72 Quarterly Journal of the Florida Academy of Sciences 

Etymology. Latin, abavus, masculine, a grandfather's grand- 
father, an ancestor. 

Description of holotype. The dorsal acetabular expansion is 
complete, moderate in size, and acute dorsally. The pit for the 
tendinous origin of the M. biceps femoris is small, round, and 
shallow. The posterodorsal border of the ilial crest slopes gently 
into the base of the dorsal acetabular expansion. The vastus prom- 
inence is much enlarged, rhomboidal in shape, strongly beveled, 
and lacks bordering ridges. The ilial crest is moderately devel- 
oped, quite thin, and its lateral surface is strongly excavated. 
Much of the anterior portion of the ilium is missing. The angle 
between the shaft and the base of the ventral acetabular expansion 
is greater than 90 degrees. Only a small part of the posterior 
border of the ventral acetabular expansion is broken. This ex- 
pansion is small and acute ventrally. The acetabulum is large and 
broadly rounded anteriorly. 

Paratypes. There is very little variation among the paratype 
ilia. One ilium has the vastus prominence not quite as strongly 
beveled, but this is probably due to individual variation. 

Sacra. Two sacra representing small leptodactylid frogs differ 
somewhat from the sacra of recent L. melanonotus. In the fossils 
the cylindrical shape of the sacral diapophyses is evident even 
though their tips are broken. The prezygapophyseal faces are 
pointed (somewhat less pointed in L. melanonotus) and have their 
long axes almost parallel to the long axis of the centrum (long axes 
more oblique in L. melanonotus). A strong thin ridge runs trans- 
versely across the top of the neural arch. The neural canal is 
much broader than high. The condyles lie very close to one an- 
other (slightly separated in L. melanonotus). The centra are broad- 
er than long through the condyles. The cotyla of both fossils are 
depressed. The width through the condyles of the F.G.S. speci- 
men is 1.6, whereas this width is 1.3 mm in the M.C.Z. specimen. 

Family Ranidae 

All New World fossils of the family Ranidae have been assigned 
to the genus Rana. Tihen (1951) and Auffenberg (1956) list Rana 
sp. from the lower Miocene of the Thomas Farm of Florida. 
Rana johnsoni La Rivers from the lower Pliocene of Storey County, 
Nevada (La Rivers, 1953), and Rana pliocenica Zweifel from the 



Holm an: Miocene Frogs 73 

middle Pliocene of Contra Costa County, California (Zweifel, 1954), 
are heretofore the earliest named species of New World Rana. 
Nine Rana species have been described from the upper Pliocene 
of Meade County, Kansas (see Taylor, 1942, and Holman, 1963b). 
These species should be considered species inquirendae until the 
Kansas bones have been re-studied. 

Pleistocene ranids definitely assigned to species are all repre- 
sented by forms living today. These fossils include: Rana cates- 
beiana (Sangamon, Meade County, Kansas, Tihen, 1954); Rana 
grylio (Illinoian?, Alachua County, Florida, Tihen, 1952), and 
Rana pipiens (various localities in Florida, Holman, 1962a; Kansas, 
Tihen, 1954; and Texas, Holman, 1962b and 1964, and Mecham, 
1959). 

Genus Rana Linnaeus 

The ilia of recent New World Rana studied (R. areolata 3, R. 
aurora 1, R. boylii 2, R. capito 1, R. cascadae 1, R. catesbeiana 17, 
R. clamitans 8, R. grylio 3, R. heckscheri 3, R. palmipes 3, R. palus- 
tris 1, R. pipiens 77, R. sylvatica 5) and of R. esculenta (1) of the 
Old World are distinct from 2 R. temporaria and a small assem- 
blage of other ranid genera available. In the former group the 
ilial crest is well developed and the vastus prominence is flattened 
laterally and is not produced above the level of the ilial crest an- 
terially. In Rana temporaria and 1 Ooeidozyga laevis the ilial 
crest is less developed, and the vastus prominence is flattened, 
but is moderately (R. temporaria) or much (O. laevis) produced 
above the level of the ilial crest anteriorly. In 1 Cacosternum 
boettzeni, 2 Mantella auriantiaca, 1 Rhacophorus buergeri, and 3 
Staurois natator the ilial crest is absent, and the vastus prominence 
is knob-like. 

New World species of Rana (including R. esculenta) separate 
into two groups based on ilial characters. In R. areolata, R. capito, 
R. cascadae, R. palmipes, R. palustris, R. pipiens, and R. sylvatica. 
the posterodorsal border of the ilial crest slopes gently into the 
dorsal acetabular expansion. In R. aurora, R. boyli, R. catesbei- 
ana, R. clamitans, R. esculenta, R. grylio, and R. heckscheri the 
posterodorsal border of the ilial crest slopes precipitously into the 
dorsal acetabular expansion. This condition was first noticed by 
Auffenberg (1956) who used different terminology to describe it. 
Whether the condition indicates natural groupings is unknown at 



74 Quarterly Journal of the Florida Academy of Sciences 



present. The ilia of Thomas Farm ranid frogs represent the genus 
Rana, being similar to the former assemblage of Rana species in 
this character. 








Fig. 2. Top left, holotype left ilium of Rana miocenica sp. nov., F.G.S. 
V-6069; top right, holotype right ilium of Rana bucella sp. nov., F.G.S. 
V-6071; bottom left, left ilium of Rana cf. R. pipiens, U.F. 10203 (each line 
represents 2 mm). Bottom right, ilium drawn from Rana pipiens to illustrate 
origin of selected muscles: A, acetabular fossa; B, tendinous origin of M. 
biceps femoris; C, fleshy origin of vastus externus head of M. triceps femoris; 
D, fleshy origin of M. gluteus; E, tendinous origin of M. sartorius; F, ten- 
dinous origin of vastus internus head of M. triceps femoris. 

Figure 2 shows the distribution of muscular origins on an ilium 
of Rana pipiens. Rana catesbeiana specimens were also dissected. 
I have attempted to use terminology consistent with the muscula- 
ture of the Rana ilium whenever possible. 

Two new species of Rana from the lower Miocene of the 
Thomas Farm of Florida are detailed in the following paragraphs. 
These fossil frogs represent the earliest ranids described from North 
America. 

Rana miocenica sp. nov. 

Holotype. Left ilium F.G.S. V-6069 (Fig. 2). From Hawthorne 
formation, lower Miocene, Arikareean; Thomas Farm, Gilchrist 
County, Florida. Collected by Pierce Brodkorb, November 14, 
1959. 



Holman: Miocene Frogs 75 

Referred element. Sacral vertebra F.G.S. V-6070. Collected 
from the same locality by the same collector in 1962. 

Diagnosis. A medium-sized Miocene Rana similar to R. areolata, 
R. capito, R. cascadae, R. palmipes, R. palustris, R. pipiens, and 
R. sylvatica in having posterodorsal border of ilial crest sloping 
gently into dorsal acetabular expansion, but differing in having 
vastus prominence relatively short, but much produced, and with 
strong bordering ridges (other species with this prominence usually 
more elongate, less produced; if bordering ridges present, never as 
extensive or strong); also differing from the above species (R. palm- 
ipes being a single exception) in having posteorodorsal border of 
ilial crest along medial attachment of dorsal fascia swollen (other 
species with this border blade-like). Measurements: greatest 
height of acetabular fossa 3.3; highest part of ilial shaft 4.0; great- 
est length of vastus prominence 3.1 mm. 

Description of holotype. Much of the dorsal acetabular ex- 
pansion is broken. The pit for the tendinous origin of the M. 
biceps femoris is moderately large and is triangular in shape. The 
posterodorsal border of the ilial crest slopes gently into the dorsal 
acetabular expansion. The vastus prominence is relatively short, 
but much produced and has strong bordering ridges. It is ovaloid 
in shape and flattened laterally. The ilial crest is well developed 
and high, and its lateral surface is moderately excavated. Part of 
the anterior portion of the ilium is broken. The angle between 
the shaft and the ventral acetabular expansion is greater than 90 
degrees. Much of the ventral acetabular expansion is broken. 
The acetabulum is moderate in size, about as long as high, and is 
moderately excavated. The anterior border of the acetabulum 
appears to be narrowly rounded, although some of the acetabular 
border is chipped. The posterodorsal border of the ilial crest along 
the medial attachment of the dorsal fascia is swollen. 

Sacrum. The referred sacrum originates from an individual 
of about the same size as the one represented by the holotype. 
The fossil vertebra is a typical Rana sacrum with one anteriorly 
directed condyle and two posteriorly directed condyles. The di- 
apophyses are cylindrical. The left diapophysis has a single an- 
terior tubercle at about the middle of its extent, and the right di- 
apophysis has a divided anterior tubercle at about the middle of 
its extent. The prezygapophyseal faces are ovaloid in shape, and 
just behind them, on the top of the neural arch, is a flattened tri- 



76 Quarterly Journal of the Florida Academy of Sciences 

angular area. The neural canal is about twice as wide as high. 
The posterior condyles are depressed and separated from each 
other by a space a little less than half the width of one condyle. 
The centrum is as broad as it is long through the condyles. The 
anterior condyle is much depressed. The width of the sacrum 
through the diapophyses is 10.0; the width of the centrum through 
the condyles is 3.2; the length of the sacrum through the con- 
dyles is 3.2 mm. 

The fact that the length of the centrum divided by the width 
of the centrum is 1.0 is interesting in that it places the fossil closer 
to R. pipiens rather than to R. catesbeiana, R. grylio, and R. heck- 
scheri based on the data of Tihen (1954, p. 219, fig. 1). 

Remarks. Rana miocenica was a frog about the size of a recent 
Rana pipiens with a snout-vent length of about 90 mm. The re- 
lationships of R. miocenica to an assemblage of recent Rana with 
the gently sloping posterodorsal border of the ilium is apparent, 
but it is difficult to ally the fossil to any one of these species be- 
cause of their great similarity to each other. Nevertheless, R. 
miocenica shows some similarity to three recent skeletones of 
R. palmipes in the development of the knob-like condition of the 
posterodorsal border of the ilial blade along the medial attach- 
ment of the dorsal fascia. In R. palmipes the condition is not as 
pronounced as in the fossil. Also, in the three recent skeletons of 
R. palmipes the sacrum is fused to the vertebra that proceeds it. 
The sacrum assigned to R. miocenica is free. 

An assemblage of ilia from the upper Pliocene of Meade 
County, Kansas are figured by Taylor (1942). Most of these are 
assigned to the genus Rana, but none are identified to species. 
These ilia have the vastus prominence elongate and without strong 
bordering ridges, thus differing from R. miocenica. Rana plio- 
cenica Zweifel from the middle Pliocene of Contra Costa County, 
California, differs from R. miocenica in that the posterodorsal bor- 
der of the ilial crest slopes precipitously into the dorsal acetabular 
expansion (Zweifel, 1954, p. 85, fig. 1). 

Rana bucella sp. nov. 

Holotype. Right ilium F.G.S. V-6071 (Fig. 2). From Haw- 
thorne formation, lower Miocene, Arikareean; Thomas Farm, Gil- 
christ County, Florida. Collected by Pierce Brodkorb, June, 1958. 



Holman: Miocene Frogs 77 

Diagnosis. A small Miocene Rana showing no close affinities 
to recent or fossil Rana species studied, and differing strikingly 
from all of these in having a very slight slope between the postero- 
dorsal border of the ilial crest and the dorsal acetabular expansion, 
and in having the pit for the tendinous origin of the M. biceps 
femoris much enlarged. Measurements: greatest height of ilial 
shaft 2.7; greatest length of biceps femoris pit 1.3 mm. 

Etymology. Latin, bucella, feminine, diminutive of buccea, 
feminine, a morsal, a mouthful, in reference to the diminutive size 
of the fossil species. 

Description of holotype. The tip of the dorsal acetabular ex- 
pansion is broken. The biceps femoris pit is much enlarged and 
is triangular in shape. The posterodorsal border of the ilial crest 
has a very slight slope into the dorsal acetabular expansion. The 
vastus prominence is relatively elongate, not highly produced, lacks 
bordering ridges, and is ovaloid in shape. The ilial crest is well 
developed and its lateral surface is rather deeply excavated. Much 
of the anterior portion of the ilium is broken. The angle between 
the ilial shaft and the ventral acetabular expansion is impossible 
to measure because so much of the ventral acetabular expansion 
is broken. The acetabulum is moderate in size, higher than long, 
and is rather shallowly excavated. The anterior border of the 
acetabulum is narrowly rounded. The posterodorsal border of the 
ilial crest along the medial attachment of the dorsal fascia is blade- 
like. 

Remarks. Rana miocenica shows certain resemblances to liv- 
ing North American species of Rana, especially in the conforma- 
tion of the posterior part of the ilium, but Rana bucella is quite 
different from all living and fossil Rana studied. A new generic 
name might be in order if it were not for the fact that other genera 
of the Ranidae such as Staurois and Ooeidozyga depart even more 
from the ilial structure of Rana. It is here suggested that R. bucella 
has no close relatives among living species of the genus. Rana 
bucella lacks the numerous small perforations of the ilium that are 
characteristic of newly metamorphosed and young ranids, thus 
the fossil represents an adult or subadult. 

Rana cf. Rana pipiens 

Material. Left ilium, U.F. 10203 (Fig. 2), right ilium, M.C.Z. 
1994. 



78 Quarterly Journal of the Florida Academy of Sciences 

Remarks. These ilia are indistinguishable from a large series 
of recent R. pipiens, and although other recent species of Rana are 
very similar to R. pipiens it seems wise to suggest that the closest 
relationships of the fossils are with this species based on present 
geographic distributions. 

M.C.Z. 1994 has already been listed as Rana sp. by Tihen 
(1951). It is a small fragmentary bone, but it has the posterodorsal 
border of the ilium sloping gently into the dorsal acetabular ex- 
pansion, and the vastus prominence indistinct and lacking border- 
ing ridges. This condition is often seen in young specimens of 
R. pipiens with ilia of about the same size as M.C.Z. 1994. 

Two other fragmentary ilia that I have been unable to locate 
(U.F. 5919) were discussed by Auffenberg (1956). He suggests 
these ilia are similar to those in an assemblage of frogs that in- 
cludes R. pipiens. 

U.F. 10203 appears to be the most complete of the four ilia 
that are similar to recent R. pipiens. A description of this bone 
is as follows. The tip of the dorsal acetabular expansion is broken. 
The pit for the origin of the M. biceps femoris is moderately large 
and is subtriangular in shape. The posterodorsal border of the 
ilial crest slopes gently into the dorsal acetabular expansion. The 
vastus prominence is relatively elongate, but rather indistinct, and 
it lacks bordering ridges. It is ovaloid in shape. The ilial crest is 
well developed and high throughout and has its lateral surface 
well excavated, especially along its dorsal portion. The anterior 
portion of the ilium is broken. The angle between the shaft and 
the ventral acetabular expansion is greater than 90 degrees. The 
ventral acetabular expansion is broken except for its base. The 
acetabulum is moderate in size, about as high as long, and is mod- 
erately excavated. The anterior border of the acetabulum is 
broadly rounded. The posterodorsal border of the ilial crest along 
the medial attachment of the dorsal fascia is blade-like. Measure- 
ments: greatest height of ilial shaft 2.9; greatest length of vastus 
prominence 2.3; greatest height of acetabulum 2.7 mm. 

Rana sp. indet. 

The following elements are assigned to the genus Rana, but 
specific allocations are not suggested: scapulae, 3 (U.F. 10204 and 
6580, M.C.Z. 3407); humeri, 7 (U.F. 10205 and 6580, M.C.Z. 2003, 
F.G.S. V-6072); radio-ulnae, 1 (U.F. 10206); tibiofibulae, 2 (M.C.Z. 
2001). 



Holm an: Miocene Frogs 79 

Family Brevicipitidae 

The only brevicipitid frog genus reported thus far from the 
North American fossil record is Gastrophryne. Auffenberg (1956) 
has listed Gastrophryne from the lower Miocene of the Thomas 
Farm of Florida; Gastrophryne carolinensis has been reported from 
the Pleistocene of Florida (Holman, 1962a); and Gastrophryne 
olivacea has been reported from the Pleistocene of Texas (Holman, 
1963a). 

Genus Gastrophryne Fitzinger 

I have re-examined the specimen (U.F. 5144) that Auffenberg 
referred to the genus Gastrophryne, and I am in full accordance 
with his generic designation. The dorsal prominence of 2 Hypo- 
pachus cuneus, and 1 H. oxyrhinus is larger and more produced 
from the shaft than that of Gastrophryne (see Holman, 1963a, p. 
156, fig. 1 for illustrations of the ilia of Hypopachus and Gastro- 
phryne). The fossil agrees with Gastrophryne in this character. 
Additional brevicipitid comparative material has become avail- 
able that indicates the fossil is indistinguishable from recent 6 
G. carolinensis, but is quite different from 3 G. olivacea. There- 
fore, I wish to tentatively refer the Thomas Farm ilium to the 
former species. 

Gastrophryne cf. Gastrophryne carolinensis 

Material. Right ilium, U.F. 5144. See Auffenberg (1956, p. 7) 
for a figure of this specimen. 

Remarks. There are striking differences between the ilia of 
G. carolinensis and G. olivacea that have been discussed by Hol- 
man (1963a, p. 155), who figures the ilia of these two species (p. 
156, fig. b and c). Although part of the ventral acetabular expan- 
sion of the fossil is broken (not as much as is shown in Auffenberg's 
figure) the fossil clearly shows its relationships are with G. caro- 
linensis. 

Discussion and Summary 

The three anuran families (Leptodactylidae, Ranidae, and Brev- 
icipitidae) detailed in this paper are all indigenous to Florida 
today. All of the three genera reported (Leptodactylus, Rana, 
and Gastrophryne) are extant, and with the exception of Lepto- 
dactylus are present in modern Florida. Of the five species dis- 



80 Quarterly Journal of the Florida Academy of Sciences 

cussed (L. abavus, R. miocenica, R. bucella, R. cf. R. pipiens, and 
G. cf. G. carolinensis), the first three are extinct, and the last two 
have been tentatively referred to species living in Florida today. 

Only one species of Leptodactylidae occurs in Florida today, 
and this form, Eleutherodactylus ricordi, has been introduced from 
the West Indies (Carr and Goin, 1955). Leptodactylus presently 
ranges from southern Texas and Sonora to Argentina, the Antilles, 
and the islands of San Andres and Providence (Smith and Taylor, 
1948). 

Six species of Rana occur in Florida today (Carr and Goin, 
1955). Three of these are large bullfrogs (R. catesbeiana, R. 
heckscheri, and R. grylio), two are moderately large (R. capito 
and R. clamitans), and one (R. pipiens) is a medium-sized frog. 
It is interesting to note that two of the fossil species (R. miocenica 
and R. cf. R. pipiens) are medium-sized frogs, whereas the third 
(R. bucella) is a diminutive Rana. The absence of large bullfrogs 
from the early Miocene of the Thomas Farm is unexplainable at 
present. 

The only brevicipitid present from the Thomas Farm shows 
its relationships with recent Gastrophryne carolinensis, a frog that 
ranges through southeastern United States today, rather than with 
the southwestern forms G. olivacea and Hypopachus. 

A more detailed discussion of the affinities of the fossil anuran 
fauna, and remarks on the paleoenvironment of the Thomas Farm 
are deferred until the remaining fossil anuran families are studied. 

Literature Cited 

Auffenberg, W. 1956. Remarks on some Miocene anurans from Florida, 
with a description of a new species of Hyla. Breviora, no. 52, pp. 
1-11, figs. 1-3. 

. 1957. A new species of Bufo from the Pliocene of Florida. Quart. 

Jour. Florida Acad. Sci., vol. 20, pp. 14-20, 2 figs. 

. 1958. A small fossil herpetofauna from Barbuda, Leeward Islands, 

with the description of a new species of Hyla. Quart. Jour. Florida 
Acad. Sci., vol. 21, pp. 248-254, 1 fig. 

. 1963. The fossil snakes of Florida. Tulane Studies Zool., vol. 10, 

pp. 131-216, 51 figs. 

Carr, A., and C. J. Goin. 1955. Guide to reptiles, amphibians, and fresh- 
water fishes of Florida. Univ. Florida Press, Gainesville, xi + 341 
pp., 30 figs., 67 pis. 



Holman: Miocene Frogs 81 

Chantell, C. J. 1964. Some Mio-Pliocene hylids from the Valentine forma- 
tion of Nebraska. American Midi. Nat., vol. 72, pp. 211-225, figs. 1-4. 

Estes, R. 1963. Early Miocene salamanders and lizards from Florida. 
Quart. Jour. Florida Acad. Sci., vol. 26, pp. 234-256, figs. 1-4. 

. 1964. Fossil vertebrates from the Cretaceous Lance formation of 

eastern Wyoming. Univ. California Publ. Geol. Sci., vol. 49, pp. 
1-180, 73 figs., 5 pis. 

Holman, J. A. 1959. Amphibians and reptiles from the Pleistocene (Illi- 
noian) of Williston, Florida. Copeia, 1959, pp. 96-102, 1 fig. 

. 1961. A new hylid genus from the lower Miocene of Florida. 

Copeia, 1961, pp. 354-355, 1 fig. 

. 1962a. Additional records of Florida Pleistocene amphibians and 

reptiles. Herpetologica, vol. 18, pp. 115-119. 

. 1962b. A Texas Pleistocene herpetofauna. Copeia, 1962, pp. 255- 

261, 1 fig. 

. 1963a. Late Pleistocene amphibians and reptiles of the Clear Creek 

and Ben Franklin local faunas of Texas. Jour. Graduate Research 
Center, Southern Methodist Univ., vol. 31, pp. 152-167, figs. 1-3. 

. 1963b. Anuran sacral fusions and the status of the Pliocene genus 

Anchylorana Taylor. Herpetologica, vol. 19, pp. 160-166, 2 figs. 

. 1963c. Reflections on two procoelous Rana catesheiana Shaw. 

Copeia, 1963, p. 558. 

. 1963d. A new rhinophrynid frog from the early Oligocene of 

Canada. Copeia, 1963, pp. 706-708, 2 figs. 

. 1964. Pleistocene amphibians and reptiles from Texas. Herpeto- 
logica, vol. 20, pp. 73-83, 4 figs. 

La Rivers, I. 1953. A lower Pliocene frog from western Nevada. Jour. 
Paleont, vol. 27, pp. 77-81, 1 fig., 1 pi. 

Lynch, J. D. 1964. Additional hylid and leptodactylid remains from the 
Pleistocene of Texas and Florida. Herpetologica, vol. 20, pp. 141- 
142. 

Mecham, J. S. 1959. Some Pleistocene amphibians and reptiles from 
Friesenhahn Cave, Texas. Southwestern Nat., vol. 3, pp. 17-27, 3 figs. 

Olsen, S. J. 1962. The Thomas Farm fossil quarry. Quart. Jour. Florida 
Acad. Sci., vol. 25, pp. 142-146. 

Smith, H. M., and E. H. Taylor. 1948. An annotated checklist and key 
to the amphibia of Mexico. Bull. U. S. Nat. Mus., no. 194, i-iv + 
118 pp. 



82 Quarterly Journal of the Florida Academy of Sciences 

Taylor, E. H. 1942. Extinct toads and frogs from the upper Pliocene 
deposits of Meade County, Kansas. Univ. Kansas Sci. Bull., vol. 28, 
pp. 199-235, 20 pis. 

Tihen, J. A. 1951. Anuran remains from the Miocene of Florida, with the 
description of a new species of Bufo. Copeia, 1951, pp. 230-235, 
2 pis. 

. 1952. Rana grylio from the Pleistocene of Florida. Herpetologica, 

vol. 8, p. 107. 

. 1954. A Kansas Pleistocene herpetofauna. Copeia, 1954, pp. 217- 

221, 2 figs. 

. 1960. Notes on late Cenozoic hylid and leptodactylid frogs from 

Kansas, Oklahoma and Texas. Southwestern Nat., vol. 5, pp. 66-70, 
6 figs. 

. 1962. A review of New World fossil bufonids. American Midi. 

Nat., vol. 68, pp. 1-50, 62 figs. 

Zweifel, R. G. 1954. A new Rana from the Pliocene of California. Copeia, 
1954, pp. 85-87, 2 figs. 

Department of Biological Sciences, Illinois State University, 
Normal, Illinois. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



FOSSIL LIZARDS FROM THE DOMINICAN REPUBLIC 
Richard Etheridge 

The known history of West Indian lizard faunas does not ex- 
tend very far into the past. The remains of lizards have been 
recovered from Indian middens and from cave deposits on a num- 
ber of islands, but even the oldest cave fossils are probably not 
older than late Pleistocene. Nevertheless, these remains clearly 
indicate that the island lizard faunas of the not-too-distant past 
were markedly different from those on the islands today. They 
include species that are now extinct, some of which were larger 
than any living member of their genus. They also include living 
species that occur on the same island today but which attained 
a greater maximum size, and living species that are now found 
only on other islands (Hecht, 1951; Etheridge, 1964 and unpub- 
lished data). 

The pattern of late Pleistocene extinction and gigantism found 
in West Indian lizards is similar to that found in the island mam- 
mals and the mammals of the North American continent, but 
quite unlike the Pleistocene history of the continental lizard faunas. 
Of the many lizard species reported from the Pleistocene of Flor- 
ida (Auffenberg, 1955, 1956; Holman, 1958, 1959a, 1959b, 1962a; 
Gut and Ray, 1963), of Oklahoma (Etheridge, 1960a), of Texas 
(Holman, 1962b, 1963), of Kansas (Etheridge, 1958, 1960b, 1961), 
and of California (Brattstrom, 1953, 1954, 1955, 1958) none of them 
is now extinct and only one of them grew to a larger or smaller 
maximum size than it does today. Undoubtedly the changing cli- 
mate of the Pleistocene greatly influenced the distribution of liz- 
ards on the continent, for some forms are known to have lived 
beyond their present range. Otherwise, at least insofar as their 
fossils indicate, North American lizards do not seem to have been 
much influence by Pleistocene events. 

The present study of lizard remains from a cave in the western 
part of the Dominican Republic contributes additional evidence 
that the lizard faunas of the West Indies, unlike those of the North 
American continent, have undergone dramatic changes in the 
not remote past. 



84 Quarterly Journal of the Florida Academy of Sciences 

Cerro de San Francisco Cave 

On August 7, 1958, Dr. Clayton E. Ray and his field party from 
the Museum of Comparative Zoology at Harvard excavated an 
exceedingly rich fossil deposit in a cave in the western part of 
the Dominican Republic. The cave lies in the top of Cerro de 
San Francisco, an isolated hill directly east of Fortaleza Pedro 
Santana in the province of San Rafael, adjacent to the Haitian bor- 
der. Cerro de San Francisco is indicated on the United States 
world aeronautical chart above the 2000 foot mark at latitude 19° 
6' N, longitude 71° 41/ W. 

According to Ray's field notes (Aug. 7-8, Aug. 31-Sept. 2, 1958) 
Cerro de San Francisco is an isolated limestone block. The cave, 
standing as it does high in this isolated hill, has no part in the 
present drainage of the land, but is clearly part of an older drain- 
age system. The cave is quite large, with at least three large 
skylights. Its floor is very dry and has many large and small 
pieces of roof debris. A pit was dug near the rear wall of the 
cave and another about 10 feet out toward the middle of the 
cave, both to a depth of about seven feet. The pit near the wall 
descended through well marked strata containing many fossils, the 
one nearer the middle of the cave descended through powdery 
white guano containing no bone. On the basis of the pit near 
the rear wall Ray divided the floor material into three bone-bear- 
ing strata as follows: 

Stratum 1. Surface to about 15 inches in depth, consisting of 
two subdivisions as follows: 

A. Upper unit, 12 inches thick. A brown earth with much 
goat dung and bone near its surface. Most of the bones were 
owl pellet components and were mainly of Rattus. 

B. Lower unit, 4-6 inches thick. A light gray, fluffy, ashy de- 
posit, very light weight and powdery, containing almost no bone. 

Stratum 2. Six to 12 inches thick. A dark brown earth with 
irregularly distributed black lenses. At least one of the lenses 
contained a potsherd and several charred sticks, and appeared to 
be a cook fire. The bones were distributed through the brown 
and black portions of the stratum but occurred primarily in a 
band about one foot thick and one foot below the surface. The 
stratum seems to be an owl deposit judging from its composition: 
Nesophontes, bats, birds, lizards, frogs, and a few very young in- 
dividuals of Isolobodon and Brotomys. The stratum also contained 
human bone fragments. 



Etheridge: Fossil Lizards 85 

Stratum 3. Three to 4 feet in thickness. A yellowish brown 
earth extending from a sharp border with the darker earth of 
Stratum 2 to a lower stratum of limestone rubble. The stratum 
contained Isolobodon, Brotomys, Plagiodontia, Nesophontes, sloth, 
bats, lizards, and birds. 

Below Stratum 3 is a layer about l-F/2 feet thick of limestone 
rubble forming a solid floor in places. Below this a layer of very 
hard, brown earth, possibly an indurated guano, extends for a 
depth of at least one foot. Neither of these two lower strata 
contained bone. 

The lizard fossils reported here are from Strata 2 and 3. Lying 
as they do below the Rattus level, they are almost certainly pre- 
Columbian in age. The presence of human bone fragments in 
Stratum 2 suggest a maximum age of not more than 4000 years, 
for according to Rouse (1964) man did not arrive in the Greater 
Antilles before 2000 B.C. Stratum 3 may be much older, but is 
probably not older than late Pleistocene. The lizard faunas of 
the two strata are identical in species composition. In spite of 
their relative youth these cave deposits contain the remains of an 
an extinct snail (Clench, 1962) and a number of extinct vertebrates. 
Among the lizard fossils is an extinct species of Leiocephalus and 
living forms of Aristelliger, Anolis, Leiocephalus, Ameiva, and 
Diploglossus. 

In the following account the minimum and maximum snout- 
vent lengths of the animals from which the fossils came were cal- 
culated by multiplying various measurements of the fossils by the 
ratio of measurements of the same elements of skeletons of modern 
specimens to their snout-vent lengths. Series of modern skeletons 
were used to diminish the error introduced by individual variation 
and ontogenetic changes in proportions. 

The type of the extinct Leiocephalus is in the Museum of Com- 
parative Zoology at Harvard. All other specimens referred to are 
in the vertebrate paleontology collections of the Florida State 
Museum. 

Gekkonidae 

Aristelliger lar Cope 1861 

Several hundred fossils of a large gecko are referred to the 
modern Hispaniolan species Aristelliger lar. The criteria used for 
their generic identification are those given by Hecht (1951) and 



86 Quarterly Journal of the Florida Academy of Sciences 

Etheridge (1964). Most of existing structural and proportional 
differences among the species of Aristelliger appear to be due 
only to differences in size. The size disparity is great; maximum 
snout-vent lengths recorded for each species are: cochranae 60 
mm, praesignis 85 mm, georgeensis 115 mm, lar 135 mm and titan 
150 mm (Hecht, 1951). 

The maximum size reached in this population was approximate- 
ly that attained by Aristelliger lar today. Calculations of snout- 
vent lengths from measurements of the smallest and largest fossils 
of each element are: based on dentaries 66-120, on maxillae 57-121, 
on parietals 89-117, on quadrates 98-120, on frontals 69-140, on 
basale 94-120, and on pelves 80-117 mm. Fusion of the parietals, 
an indication that near-adult size has been reached, does not occur 
in fossils below calculated snout-vent length of 110 mm. This pre- 
cludes the possibility that a smaller species is represented by some 
of the smaller fossils. 

The fossils exhibit an ontogenetic increase in the number of 
teeth on the dentary and maxilla (Fig. 1) similar to that demon- 
strated in other geckos, Thecadactylus rapicaudiis (Etheridge, 1964) 
Tarentola annularis and T. ephippiata (Grandison, 1961). Tooth 
counts of modern adult Aristelliger lar approximate those of fossils 
of the same size. Tooth counts of a few individuals of A. praesignis, 
A. cochranae, A. georgeensis, and A. titan, plotted on the same 
chart, indicate the possibility that A. titan may follow the same 
ontogenetic gradient as A. lar, and that the other forms may not. 
If this proves to be the case then there would appear to be no 
anatomical reason for considering the extinct Jamaican A. titan 
other than a population of A. lar grown to larger size, for they 
differ only in maximum size attained and number of teeth present 
at maximum size. 

Fossils of Aristelliger lar have also been reported from Deep 
Cave, near St. Michel de l'Atalaye, Department de l'Artibonite, 
Haiti (Hecht, 1951). This locality and the cave at Cerro de San 
Francisco lie far outside the range of the species as reported by 
Cochran (1941): the southwestern peninsula of Haiti and the Sa- 
mana Peninsula of northeastern Dominican Republic. Hecht sug- 
gests that the modern distribution of A. lar is probably relict, and 
the occurrence of fossils outside the present range indicates that at 
an earlier time the species occupied a wider range on the island 
than it does today. Dr. Albert Schwartz (in litt.) has found the 



Etheridce: Fossil Lizards 



87 



species at a number of other localities and believes that it occurs 
rather generally over the entire island. Dr. Schwartz believes 
that its apparent relict distribution is merely an artifact of collect- 
ing, and its rarity in collections is probably due to its strictly 
nocturnal habits and its preference for large Ficus and other trees 
that have prop roots and are covered with woody vines. 



31 

36 
34 

« 324 



■▲▲■ 
DAA 



oo «oe :•• • mo »o 
•OCGOO** •••<£• «w» • 

D 0«0 • OO O O 

oo o»o 



Tooth Row Length in mm. 

Fig. 1. Ontogenetic increase in the number of teeth on the dentary 
(solid symbols) and maxilla (open symbols) in Aristelliger. Small circles, 
A. lar from Cerro de San Francisco. Large circles, A. titan. Triangles, A. 
praesignis, Squares, A. cochranae expectatus. Hexagons, A. georgeensis. 



Referred Specimens. Dentaries 10051 (141), maxillae 10052 
(178), frontals 10053 (66), basale 10054 (9), parietals 10055 (13), 
quadrates 10056 (10), surangular-articular 10057 (21), pterygoids 
10058 (12), pelves 10059 (28), vertebrae 10060 (26). 



Iguanidae 

Anolis ricordii (Dumeril and Bibron) 1837 

About 80 cranial elements and eight pelves are identical in all 
major features with modern skeletons of Anolis ricordii. They 
may be distinguished from all other Hispaniolan anoles by their 



88 Quarterly Journal of the Florida Academy of Sciences 

large size and by the presence of strong rugosities on the upper 
surfaces of the skull roofing bones. 

The fossil population is calculated to have attained a maximum 
snout-vent length of about 190 mm, larger than that of any living 
species of Anolis. Measurements of the smallest and largest fos- 
sils of each element yield the following estimates of snout-vent 
length: based on dentaries 78-166, on maxillae 155-190, on frontals 
112-192, on parietals 84-141, on basale 132-167 mm. Williams 
(1960) gives the snout-vent length of adult males of A. ricordii as 
about 137 mm. He has informed me (in litt.) that the largest 
specimen in the Museum of Comparative Zoology has a snout- 
vent length of 159 mm. Thus the species has decreased about 
30 mm in maximum snout-vent length. 

Fossils of Anolis ricordii are also known from Deep Cave near 
St. Michel de l'Atalaye, Department L'Artibonite, Haiti (Hecht, 
1951). Anolis ricordii occurs over most of the island and probably 
in the vicinity of the cave today. 

Referred Specimens. Dentaries 10061 (15), maxillae 10062 (41), 
frontals 10063 (12), parietals 10064 (7), prefrontal 10065 (1), jugals 
10066 (3), pterygoids 10067 (2), postorbital 10068 (1), articular-sur- 
angulars 10069 (3), basale 10070 (4), quadrate 10071 (1), pelves 
10072 (8). 

Anolis cybotes Cope 1862 

The most abundant lizard remains are those of a moderate-size 
anole, represented by over 1000 fossils. The dentaries show a 
distinctive type of sculpturing characteristic of the modern His- 
paniolan species A. cybotes, A. armouri, A. shrevei, and A. white- 
mani. At about 40-45 mm snout-vent length, a number of closely 
spaced, shallow grooves appear on the ventrolateral face of the 
dentary. The grooves extend from the level of the last few teeth 
to the level of the dorsal process of the coronoid. Posteriorly the 
grooves are deepest and all of them end abruptly at the same level: 
anteriorly they fade out gradually. As the jaw increases in size 
the grooves deepen and become more irregular. They extend their 
coverage ventrally and anteriorly, but not posteriorly. With con- 
tinued growth the ventrolateral face of the element becomes great- 
ly swollen and the irregular grooves are thrown into a series of 
horizontal, more or less semilunar folds. At maximum size the 
sculpturing covers all of the ventral and ventrolateral face of the 



Etheridge: Fossil Lizards 89 

dentary anterior to the level of the coronoid. Posteriorly the 
swollen area ends abruptly with a heavily sculptured projection 
that has a deep recess in its posterior surface. This sequence of 
ontogenetic changes in the dentary occurs in all of the above 
mentioned species and, when they are lined up from smallest to 
largest, in an identical fashion in the fossils. The sculpturings 
that occur on the lower jaws of other species of Anolis (cristatellus, 
scriptus, gundlachi, pulchellus, krugi, and bimaculatus) have en- 
tirely different configurations. 

I am unable to distinguish the skeletons of A. cybotes, A. ar- 
mouri, A. shrevei, and A. whitemani. Anolis cybotes occurs over 
most of the island today and in the immediate vicinity of the 
cave. The other species have more restricted ranges, none of 
which includes the Cerro de San Francisco. For this reason, and 
no other, I have tentatively referred the fossils to Anolis cybotes. 

The fossil population is estimated to have reached a maximum 
snout-vent length of about 75 mm, slightly larger than that at- 
tained by most living populations. Snout- vent lengths calculated 
for the smallest and largest fossils of each element are: based on 
dentaries 42-75, on parietals 55-75, on quadrates 58-71, and on 
basale 58-69 mm. Williams (1960) gives the snout-vent length of 
adult males of A. cybotes as 67 mm. He has informed me (in lift.) 
that the race A. cybotes haetianus from the southwestern peninsula 
of Haiti reaches a maximum of 79 mm, but that over the rest of 
the island the maximum snout-vent length attained is 70 mm 
(measurements based on specimens in the Museum of Comparative 
Zoology). 

Referred Specimens. Dentaries 10073 (377), maxillae 10074 
(506), postorbitals 10075 (6), frontals 10076 (82), parietals 10077 
(28), pterygoids 10078 (2), quadrates 10079 (6), premaxillae 10080 
(2), prefrontals 10081 (2), articular-surangulars 10082 (32), jugals 
10083 (11), basale 10084 (39), interclavicle 10085 (1), pelves 10086 
(44). 

Anolis chlorocyanus Dumeril and Bibron 1837 

A single dentary is referred to Anolis chlorocyanus, primarily 
on the basis of its slender form. The jaw measures 9.4 mm from 
the symphysis to the posterior border of the last tooth, 1.3 mm 
deep at the position of the last tooth and 0.9 mm deep midway 
between the symphysis and the last tooth. Using these three 



90 



Quarterly Journal of the Florida Academy of Sciences 



measurements the fossil has been compared with dentaries of other 
Hispaniolan anoles by discriminant function analysis programmed 
for a 1620 IBM computer (Rao, 1958, p. 239). The species number 
of dentaries compared with the fossil are A. coelestinus 3, distichus 
4, cybotes 8, aliniger 2, singularis 2, ricordii 2, chlorocyanus 4, 
whitemani 4, monticola 2, cochranae 1, cristophei 2, etheridgei 2, 
koopmani 2, shrevei 2, semilineatus 2, hendersoni 2, and olssoni 4. 
The analysis shows that the dentaries of hendersoni and singularis 
are relatively more slender than the fossil, the dentary of chlor- 
ocyanus has the same shape as the fossil, and the dentaries of all 
other species are relatively less slender. Measurements of the 
dentary of a modern individual of A. chlorocyanus 64 mm snout- 
vent length are very close to those of the fossil: tooth row length 
9.4, depth at the last tooth 1.4, and depth midway between the 
first and last tooth 1.0 mm. The jaw contains 26 teeth, the fossil 
contained 28. The fossil is therefore believed to have come from 
an individual of Anolis chlorocyanus about 64 mm snout-vent 
length. The species occurs over most of the island and probably 
lives in the immediate vicinity of Cerro de San Francisco today. 
Referred Specimen. Dentary 10087. 

Leiocephalus Gray 

Until recently the genus Leiocephalus included a number of 
South American species as well as those in the West Indies. The 
continental forms have been shown to be unrelated to the insular 
species, and Leiocephalus has been restricted to the latter (Ether- 
idge, 1965). Leiocephalus now occupies the islands of Hispaniola, 
Navassa, the Caymans, Cuba, and the Bahamas, and a recently 
extinct form occurred on Martinique. An extinct species has been 
described from the late Pleistocene of Barbuda and other extinct 
forms have been reported, but not described, from the Pleistocene 
of Hispaniola and Jamaica (Etheridge, 1964) and the Miocene of 
Florida (Estes, 1963). On the basis of osteological characteristics 
of Leiocephalus given in these publications a number of fossils 
from the cave in Cerro de San Francisco may be referred to the 
genus. Two species are represented in the fossils, a small one 
assigned to a modern Hispaniolan species and a very large one 
that is extinct. In reference to its most obvious diagnostic char- 
acter, an open Meckelian groove, the extinct species may be 
known as 



Etheridge: Fossil Lizards 91 

Leiocephalus apertosulcus, new species 

Holotype. A right dentary, No. 3404 in the vertebrate paleon- 
tological collections of the Museum of Comparative Zoology, Har- 
vard University, Cambridge, Mass. 

Type Locality. Stratum 2, cave in Cerro de San Francisco, 
Municipio Pedro Santana, Provincia San Rafael, Republica Do- 
minica. Probably Late Pleistocene. Collected by Dr. Clayton 
E. Ray in August or September, 1958. 

Referred Specimens. Dentaries 10088-9 (50), maxillae 10090-91 
(36), frontals 10092 (6), parietals 10093 (6), jugals 10094 (3), post- 
orbital 10095, pterygoid 10096, quadrates 10097 (2), articular + 
surangular 10098, prefrontal 10098, basale 10099, pelves 10100 
(15), caudal vertebrae 10101 (6). 

Diagnosis. Leiocephalus apertosulcus differs from all other 
species in the genus, both living and extinct, in having Meckel's 
canal exposed as a deep groove along the lingual face of the den- 
tary; in all other forms it is a tubular cavity completely surrounded 
by bone. The maximum snout-vent length of this species was at 
least 150 mm and possibly as great as 200 mm. This exceeds the 
maximum length attained by any living species (130 mm) and is 
equaled only by the extinct Barbudan species Leiocephalus cuneus. 




Fig. 2. Lingual view of the dentary of Leiocephalus apertosulcus; type 
specimen. 

Description of Type (Fig. 2). The specimen is 17.2 mm long 
from its broken posterior border to the symphysis. The tooth row, 
measured in a straight line from the projected anterior border of 
the first tooth to the projected posterior border of the last tooth 
(both are missing) is 16.7 mm long. The height at the position 
of the penultimate tooth is 4.0 mm. There are 23 teeth or unoc- 
cupied alveoli; missing are (front to back) teeth number 1, 2, 4, 
18, find 23, and tooth number 13 lacks its entire crown. Teeth 



92 Quarterly Journal of the Florida Academy of Sciences 

number 3, 5, and 6 are simple, bluntly conical and curve inward 
and slightly backward. Tooth number 9 and all teeth posterior 
to it have a tall, slender, straight-sided shaft and a crown that is 
flared in an anterior-posterior direction, linguo-labially compressed 
to form a moderately sharp cutting edge, strongly tricuspid and 
curved inward. The anterior and posterior cusps of each tooth 
are smaller than the median cusp and separated from it by a wide 
groove that fades out at the base of the crown. The occlusal edge 
of each of these teeth is slightly oblique to the main axis of the 
tooth row. The tricuspid teeth become closely crowded pos- 
teriorly, where the anterior cusp of each tooth is overlapped labially 
by the posterior cusp of the preceding tooth. About 40 per cent 
of each tooth rises above the alveolar border of the dentary. Teeth 
number 7, 8, and 9 are transitional in form between the anterior 
simple teeth and the posterior tricuspid ones. A vertical row of 
very small foramina penetrate the lingual face of the dentary in 
the narrow spaces between the teeth. 

The labial face of the dentary is smooth and convex Five large 
mental foramina form a row on the labial face between the third 
and tenth tooth; a smaller foramen is present below this row at 
the position of the eighth tooth. A large, shallow, triangular and 
slightly concave depression in the posterodorsal part of the labial 
face extends to the level of the twentieth tooth, marking the for- 
mer position of the anterolateral process of the coronoid. The 
lingual face of the dentary is produced medially as a narrow shelf 
below the base of the tooth row from the symphisis to the nine- 
teenth tooth; beyond this the shelf has been broken away. Meck- 
el's canal is exposed lingually as a wide, open groove that extends 
the entire length of the dentary. 

Referred Specimens: Dentaries. In addition to the type there 
are 28 nearly complete dentaries and fragments of 22 others. In 
the complete dentaries the length of the tooth row, measured as 
in the type, ranges from 9.4-18.2 mm. The number of teeth in- 
creases ontogenetically from 17 in the smallest to 25 in the four 
largest dentaries (Fig. 3). The first two teeth are present in sev- 
eral specimens and, like the third, fifth, and sixth teeth of the 
type are simply pointed. The first tooth with a distinctly flared 
and tricuspid crown (number 9 in the type) varies from number 
7 to 11. There are 4-7 mental foramina. They occur as far for- 
ward as the position of the first tooth and as far posteriorly as the 



Etheridge: Fossil Lizards 93 

position of the sixteenth. In dentaries larger than the type the 
upper half of the labial face becomes flattened to slightly convex 
posteriorly and develops light surface rugosities. Pathological mal- 
formations of bone are present in two of the dentaries. In all other 
respects the additional dentaries are very similar to the type. 
Particularly important is the consistant presence of an open Meck- 
elian groove. In one specimen the upper and lower borders of 
the groove converge and touch for a short distance below the 
twelfth to fourteenth tooth, but in all others the borders of the 
groove are widely separated for their entire length. 



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Tooth Row Length in mm. 

Fig. 3. Ontogenetic increase in the number of teeth on the dentary 
(solid symbols) and maxilla (open symbols) in two species of Leiocephalus. 
Circles, L. apertosulcus. Squares, L. personatus from Cerro de San Francisco. 
Triangles, modern L. personatus. 

Maxillae. Ten nearly complete maxillae and fragments of 26 
others. The length of the tooth row varies from 11.7-17.0 mm. 
The number of teeth increases ontogenetically from 17-23 (Fig. 3). 
The crown profiles of the teeth are similar to those of the den- 
tary: simply pointed anteriorly and tricusped posteriorly. The 
first tooth with a distinctly flared, tricuspid crown varies from 
number 5 to 8. The addition of one half of the number of pre- 
maxillary teeth (3-4 based on living species) to those of each max- 
illa places the transition from simple to tricuspid at approximately 
the same position that it occurs on the dentary. From 6-9 mental 
foramina form an irregular row above the levels of the second to 
fourteenth tooth. From 3-5 additional foramina penetrate the 
ascending nasal process above the row of labial foramina. 

Frontals. Six frontal bones lack parts of their nasal processes 



94 Quarterly Journal of the Florida Academy of Sciences 

but are otherwise nearly complete. The largest measures 4.1 mm 
across its narrowest interorbital width and 12.3 mm wide along 
its parietal border. The dorsal surface is concave between the 
orbits, flattening out toward the nasal-prefrontal and parietal bor- 
ders. A smooth, shallow, semilunar depression in the upper me- 
dial face of each nasal process indicates the former position of 
the nasal bone and a depression along the ventrolateral face of 
each nasal process indicate the former position of the prefrontal. 
These sutural surfaces are not continuous across the upper face 
of the nasal processes, indicating that the nasals and prefrontals 
were separated posteriorly above by the anteriorly projecting nasal 
processes of the frontal. In the middle of the parietal border a 
deep, U-shaped notch, bordered on each side by a rounded pro- 
jection, forms the anterior border of the parietal foramen. The 
entire upper surface of the frontal is moderately rugose; however, 
the rugosities do not form regularly arranged areas correspond- 
ing to the scales that were above them. 

Parietals. Six parietals exhibit the ontogenetic change in the 
shape of the roofing part that is characteristic of all of the larger 
members of the genus (Etheridge, 1965). The largest measures 
13.7 mm wide across its frontal border and 7.3 mm from the frontal 
border to the occipital crest. The smallest is 9.6 mm wide across 
its frontal border. The upper surface of the roofing part is mod- 
erately rugose in the larger fossils; however, the rugosities do not 
form regularly arranged areas corresponding to the scales that 
were above them. 

Jugals. The largest of three jugals lacks the distal extremity 
of its temporal process, but the estimated straight-line distance 
from the tip of the maxillary process to the tip of the temporal 
process is 15 mm. One (in two specimens) or two large suborbital 
foramina penetrate the labial face of the proximal part of the max- 
illary processes; three smaller foramina penetrate the ventral edge 
of the temporal process. 

Postorbital A single postorbital is 7.4 mm high from the top 
of its frontal process to the ventral border. The outer face of the 
element is strongly sculptured. 

Pterygoid. A left pterygoid is 7.4 mm wide between the pos- 
teromedial corner of the palatine portion and the lateral extremity 
of the ectopterygoid process. The anterior end of the palatine 
process and the posterior end of the quadrate process are missing. 



Etheridge: Fossil Lizards 95 

The medial part of the ventral surface of the palatine process is 
slightly sculptured, but there is no trace of teeth or unoccupied 
alveoli. 

Quadrates. The larger of two quadrates measures 5.8 mm from 
the top of the cephalic condyle to the bottom of the ventral con- 
dyle. 

Articular + surangular. A fused articular and surangular is 
19.3 mm long; its articular part is 16.1 mm long. The former 
positions of the angular, coronoid, and dentary are clearly indi- 
cated by sutural scars. The angular extended posteriorly below 
the ventral faces of the articular and surangular almost to the 
level of the articular condyle. The dentary extended at least as 
far as the posterior limit of the coronoid. A short, robust angular 
process projects medially from the articular condyle for a distance 
of 2.1 mm. Its dorsal surface is smoothly convex, becoming some- 
what flattened and turned upward distally. The convex anterior 
border of the angular process is continuous with the medial border 
of the articular. The retroarticular process projects posteriorly 
from the articular condyle for a distance of 3.2 mm. The lateral 
and medial borders are somewhat raised. The angular and retro- 
articular processes are united posteromedially by a thin shelf 
whose margin forms a nearly straight line between the distal ex- 
tremities of the two processes. 

Prefrontal. A single prefrontal is 8.7 mm long from the an- 
terior tip of the maxillary process to the posterior tip of the frontal 
process. Its outer surfaces are distinctly rugose. 

Basale. A basale, consisting of a basisphenoid, two otic ele- 
ments, and the occipital bones, is apparently from a subadult in- 
dividual. The bones were held together by cave earth that filled 
the brain cavity, but they fell apart when the matrix was removed. 
The distance between the extremities of the basicranial tubercles 
is 5.0 mm. 

Pelves. Fifteen pelves are apparently from adult individuals, 
for in all of them the ischium, ilium, and pubis are firmly ankylosed. 
Measurements of a complete specimen are: greatest length from 
the anterior end of the pubis to the posterior end of the ilium 24.7, 
greatest height from the ventral border of the ischium to the upper 
border of the posterior prominence of the ilium 13.4, greatest di- 
ameter of the acetabulum 5.0 mm. The acetabular diameter of 
the smallest specimen is 4.3, of the largest 6.4 mm. 



96 Quarterly Journal of the Florida Academy of Sciences 

Caudal Vertebrae. Three anterior segments and three pos- 
terior segments of autotomic caudal vertebrae are referred to this 
species. The diameter of the neural arch just anterior to the con- 
dyle of the posterior segments ranges from 1.7-2.2 mm. Eight 
smaller posterior segments may be from subadults or from a more 
posterior part of the tail of this species, or they may have come 
from individuals of the smaller species of Leiocephalus in the cave. 
In all of the posterior segments a thin, median crest extends along 
the top of the neural arch from the neural spine to the fracture 
plane, where it rises to form the posterior part of what was probably 
a narrow, triangular spine above the fracture plane prior to dis- 
sociation of the two segments. The anterior border of the pos- 
terior segment (the posterior border of the fracture plane) is more 
or less vertical except for a notch on each side near the bases of 
the neural arch, the notch marking the former position of the base 
of the transverse process of the anterior segment. 

The anterior segments of the autotomic caudal vertebrae also 
possess a thin, median crest that rises steeply as it approaches the 
fracture plane. Their transverse processes are spatulate, slightly 
tapered and oriented more or less laterally. 

Comparisons. The presence of an open Meckelian groove in 
the dentary of Leiocephalus apertosulcus is unique within the 
genus. In all other forms, both living and extinct, Meckel's canal 
is a tubular cavity completely surrounded by bone. Among the 
iguanid genera allied to Leiocephalus (Tropidurus, Liolaemus, 
Stenocercus, Proctotretus, Platynotus, Plica, Ophryoessoides, Uran- 
oscodon, and Urocentron) an open Meckelian groove occurs only 
in some species of Liolaemus (Etheridge, 1965). Leiocephalus and 
Liolaemus show many similarities; present in both but absent in 
all other genera listed above are a large descending process of the 
coronoid that laterally overlaps the dentary, evident by the coro- 
noid scar on the fossil dentaries, and a lamination of the nasal 
spine of the premaxilla by the nasal bones. On the basis of these 
and other similarities of the skeleton and integument I suggested 
that, of the genera listed above, Liolaemus is probably most closely 
allied to Leiocephalus (Etheridge, 1965). There are, however, nu- 
merous osteological differences between the two, some of which 
are evident in the fossils and permit their certain identification as 
Leiocephalus: the presence of vertical rows of small foramina in the 
lingual face of the jaws in the narrow spaces between the teeth, 



Ethetudge: Fossil Lizards 97 

the strongly converging lateral borders of the adult parietal, and 
the presence of a vertical spine-like process above the plane of 
fracture of the autotomic caudal vertebrae. Although the His- 
paniolan fossils are clearly referrable to the genus Leiocephalus, 
the presence of an open Meckelian canal in this extinct species 
provides additional evidence for the close relationship of Leio- 
cephalus with Liolaemus. 

The snout-vent lengths of the animals from which the fossils 
came have been calculated by multiplying various measurements 
of the same elements of living species to their snout-vent lengths. 
Assuming the proportions that existed in the extinct species fall 
within the limits of those which exist among all of the living forms, 
a minimum and maximum estimate of snout-vent length may be 
obtained for the extinct species. The minimum and maximum esti- 
mates for the largest fossil of each of several elements are as fol- 
lows: dentary 142-173 mm, maxilla 144-178 mm, parietal 141-178 
mm. frontal 139-161 mm, postorbital 133-185 mm, articular + 
surangular 147-192 mm, and pelvis 161-202 mm. The maximum 
snout-vent length attained by any living species is 130 mm re- 
corded for Leiocephalus carinatus microcyon (Schwartz, 1959). The 
largest living Hispaniolan species is L. melanochloris, which attains 
a maximum snout-vent length of 108 mm (Cochran, 1941). All of 
the minimum values calculated for Leiocephalus apertosulcus ex- 
ceed 130 mm; thus, the species probably attained a greater size 
than any living species of the genus, and certainly must have 
grown larger than any living Hispaniolan form. On the basis of 
the same type of calculations the maximum snout-vent length at- 
tained by the extinct Barbudan species, L. cuneus, was also esti- 
mated to have been nearly 200 mm (Etheridge, 1964). 

Leiocephalus personatus Cope 1862 

The smaller of the two species of Leiocephalus from the cave 
in Cerro de San Francisco appears to be referable to the extant 
Hispaniolan species, L. personatus. This species and L. pratensis 
differ from all other forms of Leiocephalus in that the upper sur- 
faces of the frontal and parietal bones are distinctly sculptured 
after the pattern of scales that were above them. Impressions of 
the posterior parts of the interparietal and the inner parietals and 
of the medial parts of the outer parietal scales are present on the 



98 Quarterly Journal of the Florida Academy of Sciences 

roof of the parietal bone, and impressions of the anterior parts 
of the interparietal and inner parietals and of the frontoparietals 
and posterior frontal scales are present on the roof of the frontal 
bone. These scale impressions are clearly evident on the fossil 
parietals and frontals of the smaller species. The skull of Leio- 
cephalns pratensis differs from that of L. personatus in that the 
interorbital width of the frontal bone is relatively greater: the 
ratio of the posterior width of the frontal to the narrowest inter- 
orbital width is about 2.3 in pratensis and about 3.3 in personatus. 
This ratio is 3.5 in the fossil frontals. 

Fossil maxillae and dentaries referred to L. personatus lack 
the open Meckelian groove of L. apertosulcus, excluding the possi- 
bility that these small jaws are from subadult individuals of the 
latter. Furthermore, a plot of the maxillary and dentary tooth row 
lengths against the number of teeth indicates that different gradi- 
ents in the ontogenetic increase in tooth numbers exist between 
the two species (Fig. 3). In this and in all other respects the fos- 
sil maxillae and dentaries are indistinguishable from those of Re- 
cent L. personatus. 

Measurements of the largest fossils are as follows: dentary tooth 
row 8.6, maxillary tooth row 7.9, anterior width of parietal roof 
7.1, posterior width of frontal 4.8 mm. Using measurements of 
Recent skeletons of L. personatus as a basis, the snout-vent lengths 
calculated for the largest fossils are: based on dentary 75 mm, on 
maxilla 74 mm, on parietal 73 mm, on frontal 67 mm. These esti- 
mates are within the range of maximum snout-vent lengths of the 
various living races of the species: 55 mm in L. p. louisae to 83 
mm in L. p. mentalis (Cochran, 1941). 

Leiocephalus personatus occurs throughout the island today. 

Referred Specimens. Dentaries 10102 (10), maxillae 10103 (3), 
frontals 10104 (2), parietals 10105 (4). 

Teiidae 

Ameiva chrysohema Cope 1869 

Several cranial elements, pelves, and vertebrae are referred to 
Ameiva chrysohema because of their large size and close structural 
resemblance to modern skeletons of the species. The jaws of 
A. chysolaema differ from those of A. taeniura and A. lineohta 
by the possession of no more than 4 tricuspid teeth on the rear of 



Etheridge: Fossil Lizards 99 

the maxilla and no more than 5 tricuspid teeth on the rear of the 
dentary. A. taeniura and A. lineolata have from 6-11 tricuspid 
teeth on the rear of the maxilla and from 7-13 on the rear of the 
dentary. The fossil jaws assigned to this species have from 0-3 
tricuspid teeth. 

Ameiva chrysolaema reaches a maximum snout-vent length of 
160 mm, A. lineolata a maximum of 56 mm, and A. taeniura a max- 
imum of 80 mm (Schwartz, in litt.). Snout-vent lengths of the 
smallest and largest fossils referred to A. chrysolaema are: based 
on dentaries 97-125 mm, on maxillae 105-114 mm, on frontals 104- 
128 mm, on quadrate 134 mm, and on pelves 90-125 mm. Thus 
even the smallest fossil exceeds A. taeniura and A. lineolata in size. 

Three ameivas occur on Hispaniola today. A. chrysolaema and 
A. taeniura are widely distributed and probably occur in the im- 
mediate vicinity of Cerro de San Francisco. A. lineolata is re- 
stricted to certain limited areas and probably does not live near 
the cave today. Two other species are confined to off-shore is- 
lands: A. barbouri to Goanve Island and A. rosamonde to Saona 
Island. 

Referred Specimens. Dentaries 10106 (9), maxillae 10107 (2), 
frontals 10108 (2), quadrate 10109 (1), pelves 10110 (4), vertebrae 
10111 (6). 

Ameiva taeniura Cope 1862 

Several jaws and a pelvis are referred to Ameiva taeniura be- 
cause of their moderate size and possession of at least 7 tricuspid 
teeth on the rear of the jaws. Although I am unable to distinguish 
structurally the modern skeletons of A. taeniura and A. lineolata 
the fossils are too large to be assigned to A. lineolata. Snout-vent 
length estimates for the fossils are: based on dentaries 76-81 mm, 
on maxillae 70-76 mm, on pelvis 72 mm. 

Referred Specimens. Dentaries 10112 (3), maxillae 10113 (3), 
pelvis 10114 (1). 

Anguidae 

Diploglossus sternurus Cope 1862 

About 100 fossils are referred to Diploglossus sternurus entirely 
on the basis of their large size. The six diploglossine lizards known 
to occur on Hispaniola today, Diploglossus curtissi, D. costatus, D. 
sternurus, D. darlingtoni, Wetmorina haetiana and Saurisia sep- 



100 Quarterly Journal of the Florida Academy of Sciences 

soides, are very similar osteologically; they differ primarily in skull 
and girdle proportions and in maximum size attained. The maxi- 
mum snout-vent length in these forms is as follows: D. sternurus 
230 mm, D. costatus 127 mm, D. curtissi 86 mm, D. darlingtoni 67 
mm, W. haetiana 87 mm, and S. sepsoides 46 mm (Cochran, 1941; 
Schwartz, unpublished manuscript). The fossil basale and pelves 
in which the components are completely ankylosed indicate that 
two size classes of Diploglossus are present, one with a maximum 
snout-vent length of 120-130 mm and one with a maximum length 
of 210-250 mm. These figures are close to the maximum sizes of 
D. costatus and D. sternurus today. For this reason all fossils esti- 
mated to have come from animals over 130 mm snout-vent length 
are referred to Diploglossus sternurus. This species occurs widely 
over the island and probably lives in the immediate vicinity of the 
cave today. 

Referred Specimens. Dentaries 10115 (38), maxillae 10116 (22), 
premaxillae 10117 (4), parietals 10118 (4), articulars 10119 (5), ptery- 
goids 10120 (4), basale 10121 (1), pelves 10122 (3), sacral vertebrae 
10123 (5), caudal vertebrae 10124-5 (11). 

Diploglossus costatus Cope 1861 

The fossil pelves and basale from adult individuals calculated 
to have been 120-130 mm snout-vent length are referred to this 
species. About 100 other fossils of Diploglossus are estimated to 
have come from animals less than 130 mm snout-vent length. 
There is no way to determine whether fossils of this size are from 
D. costatus or from subadult D. sternurus. Both species are prob- 
ably represented in these smaller fossils. Diploglossus costatus 
also occurs widely over the island and probably in the immedi- 
ate vicinity of the cave today. The ranges of the other Hispaniolan 
diploglossines do not now include Cerro de San Francisco. 

Referred Specimens. Pelves 10126 (3), basale 10127 (2). Re- 
ferred to Diploglossus sp.: dentaries 10128 (34), maxillae 10129 (31), 
articulars 10130 (4), pterygoids 10131 (2), sacral vertebrae 10132 
(4), caudal vertebrae 10133-4 (25). 

Absence of Anolis disttchus 

Anolis distichus is widespread on Hispaniola and has been 
taken in the immediate vicinity of Cerro de San Francisco. If it 



Etheridce: Fossil Lizards 101 

had lived there at the time the cave deposits were formed it might 
reasonably be expected among the fossils. The argument for this 
concerns the manner in which the deposits were formed. Nearly 
all of the fossiliferous West Indian caves contain abundant chirop- 
teran remains. Most of the other vertebrate fossils appear to 
have originated from owl pellets (Miller, 1929; Anthony, 1919; 
Hecht, 1951). This is almost certainly the origin of the lizard 
fossils in the cave at Cerro de San Francisco. The remains of a 
barn owl, Tyto alba, a burrowing owl, Speotyto cunicularia, and one 
or two other owls, probably of the genus Asio, have been identified 
from this cave (Brodkorb, in litt.). The extinct, giant barn owl, 
Tyto ostologa, described from a cave in Haiti (Wetmore, 1922) is 
absent. Hecht (1951) reported the predalion by barn owls on 
Anolis, Diploglossus and Aristelliger in Jamaica. Wetmore and 
Swales (1931) reported the remains of Anolis ricordii and Ameiva 
sp. in modern pellets of Tyto alba on Hispaniola. I have examined 
modern owl pellets from near the mouth of a cave near Boca de 
Yuma, Province Altagracia, Dominican Republic, collected by Clay- 
ton Ray. They contain a large number of bones of Anolis ricordii, 
A. chlorocyanus, A. distichus, and A. cybotes. A single intact pel- 
let contained bones of at least 16 individuals of Anolis. 

Barn owls rarely hunt before dark. Arboreal, nocturnal lizards, 
such as Aristelliger, and lizards that sleep in exposed places, such 
as Anolis, are to be expected in their pellets. It is difficult to im- 
agine what sort of opportunity they might have to prey on strictly 
diurnal lizards that sleep in concealed places, such as Leiocephalus 
or Ameiva. The more diurnal, burrowing owls may be the chief 
predators of the latter. As one might expect, fossorial lizards 
(Amphisbaena) and very small lizards (Sphaerodactylus) do not 
occur in the cave deposits. 

Except for Leiocephalus apertosulcus, which is extinct, all of 
the lizards identified among the Cerro de San Francisco fossils 
probably occur in the immediate vicinity of the cave today. Other 
lizards that probably occur there now are Amphisbaena manni, 
Sphaerodactylus sp., Anolis olssoni and A. distichus. Amphisbaena 
and Sphaerodactylus have not been found in any West Indian cave 
deposits and are not to be expected here. Anolis olssoni, because 
of its small size and habitat of low bushes and grass is probably 
also not to be expected. In the owl pellets from Boca de Yuma 
the only local anole not present in the pellets is Anolis semilineatus, 



102 Quarterly Journal of the Florida Academy of Sciences 

a species with similar habits and of similar size to A. olssoni. 
Thus, the only lizard that occurs at Cerro de San Francisco today 
but not in the cave, whose presence among the fossils may be 
reasonably expected if it had lived there, is Anolis distichus. The 
minimum size of fossil Anolis in the cave is 35 mm snout-vent 
length, well below the maximum size of 50 mm of A. distichus. 
That this species is subject to owl predation is verified by its 
presence in large numbers in the modern pellets from Boca de 
Yuma. It is difficult to account for the absence of Anolis distichus 
in more than 2000 lizard fossils except by the assumption that it 
actually did not occur there when the deposits were formed. 

Summary 

Many thousands of vertebrate fossils were recovered from a 
cave in Cerro de San Francisco in the western part of Dominican 
Republic. Most of the non-chiropteran remains are probably from 
owl pellets. Among the fossils are eleven species of lizards, one 
of which is extinct; the others probably occur in the vicinity of 
the cave today. 

Bones of a large gekko, Aristelliger lar, are from individuals be- 
tween 57-140 mm snout-vent length. The largest slightly exceed 
the maximum size of the species today. 

A large anole, Anolis ricordii, is represented by bones from in- 
dividuals between 78-192 mm snout- vent length. The largest ex- 
ceed by 30 mm the maximum size of the species today. Fossils 
of Anolis cybotes are from individuals between 42-75 mm snout- 
vent length, and a single bone of Anolis chlorocyanus is from an 
individual 64 mm snout-vent length. Both species attained ap- 
proximately the same maximum size as they do today. 

Two anoles, Anolis olssoni and Anolis distichus, that occur near 
the cave today were not identified among the fossils. The absence 
of Anolis distichus probably indicates that it did not actually live 
there at the time the cave deposits were formed. 

Two species of Leiocephalus were recovered from the cave. 
One of them, L. apertosulcus, is extinct. It is not closely related 
to any living form and differs from all of them in possessing an 
open Meckelian groove. It is calculated to have reached a max- 
imum snout-vent length of about 200 mm, larger than any living 
member of the genus and equaled in size only by the extinct Bar- 



Etheridge: Fossil Lizards 103 

budan species L. cuneus. The smaller species is referred to Leio- 
cephalus personatus. It reached a maximum snout-vent length of 
about 75 mm, within the range of maximum sizes attained by the 
various races of that species today. 

Two ameivas are identified, Ameiva chrysolaema from individ- 
uals between 90-134 mm snout-vent length and Ameiva taeniura 
from individuals between 70-81 mm snout-vent length. Neither 
species exceeded the maximum size attained by individuals today. 

Two galliwasps, Diploglossus sternurus and D. costatus, are 
present among the fossils. The largest specimens of D. sternurus 
are from individuals 210-250 mm snout-vent length, and the largest 
specimens of D. costatus from individuals 120-130 mm snout-vent 
length, approximately the same maximum size reached by these 
species today. 

Acknowledgments 

The material reported here was collected by Dr. Clayton E. 
Ray, Dr. A. Stanley Rand, and Prof. Eugenio de Jesus Marcano 
F. with financial assistance from a Sigma Xi-RESA grant-in-aid and 
from the Museum of Comparative Zoology at Harvard. Financial 
support for the preparation of specimens came from a National 
Science Foundation grant GB-178. I am grateful to Dr. J. C. Dick- 
inson, Jr., Director of the Florida State Museum, and Dr. Clayton 
E. Ray, formerly Assistant Curator of Natural Sciences, for pro- 
viding me with facilities at the museum and financial support dur- 
ing the summer of 1963, when this study was begun. I am also 
grateful to Dr. Ernest E. Williams of the Museum of Comparative 
Zoology at Harvard and to Dr. Charles Walker of the University 
of Michigan Museum of Zoology for the loan of skeletons. Cor- 
respondence with Dr. Williams and with Dr. Pierce Brodkorb of 
the University of Florida and Dr. Albert Schwartz of Miami, 
Florida, on various aspects of this study has been extremely help- 
ful. 

Literature Cited 

Anthony, Harold E. 1919. Mammals collected in eastern Cuba in 1917, 
with descriptions of two new species. Bull. Amer. Mus. Nat. Hist., 
vol. 41, pp. 625-643. 

Auffenberg, Walter. 1955. Glass lizards (Ophisaurus) in the Pleistocene 
and Pliocene of Florida. Herpetologica, vol. 11, pp. 133-136. 



104 Quarterly Journal of the Florida Academy of Sciences 

. 1956. Additional records of Pleistocene lizards from Florida. Quart. 

Jour. Florida Acad. Sci., vol. 19, pp. 157-167. 

Brattstrom, Bayard H. 1953. Records of Pleistocene reptiles from Cali- 
fornia. Copeia, no. 3, pp. 174-179. 

. 1954. Amphibians and reptiles from Gypsum Cave, Nevada. Bull. 

So. California Acad. Sci., vol. 53, pp. 8-12. 

. 1955. A small herpetofauna from the Pleistocene of Carpinteria, 

California. Copeia, no. 2, pp. 138-139. 

. 1958. New records of Cenozoic amphibians and reptiles from 

California. Bull. So. California Acad. Sci., vol. 57, pp. 5-12. 

Clench, William J. 1962. New species of land mollusks from Republica 
Dominica. Breviora, no. 173, pp. 1-5. 

Cochran, Doris M. 1941. The herpetology of Hispaniola. Bull. U. S. 
Nat. Mus., no. 177, vii + 398 pp. 

Estes, Richard. 1963. Early Miocene salamanders and lizards from Flor- 
ida. Quart. Jour. Florida Acad. Sci., vol. 26, no. 3, pp. 234-256. 

Etheridge, Richard. 1958. Pleistocene lizards of the Cragin Quarry Fauna 
of Meade County, Kansas. Copeia, no. 2, pp. 94-101. 

. 1960a. Additional notes on the lizards of the Cragin Quarry 

Fauna. Papers Michigan Acad. Sci., Arts and Lett., vol. 45, pp. 
113-117. 

. 1960b. The slender glass lizard, (Ophisaurus attenuatus, from the 

Pleistocene (Illinoian Glacial) of Oklahoma. Copeia, no. 1, pp. 46-47. 

. 1961. Late Cenozoic glass lizards (Ophisaurus) from the southern 

Great Plains. Herpetologica, vol. 17, no. 3, pp. 179-186. 

. 1964. Late Pleistocene lizards from Barbuda, British West Indies. 

Bull. Florida State Mus. ; vol. 9, no. 2, pp. 43-75. 

. 1965. The systematic status of West Indian and South American 

lizards referred to the iguanid lizard genus Leiocephalus. Copeia, in 
press. 

Grandison, Alice G. C. 1961. Preliminary notes on the taxonomy of 
Tareniola annularis and T. ephippiata (Sauna: Gekkonidae). Zool. 
Meded., vol. 38, no. 1. pp. 1-13. 

Gut, H. James, and Clayton E. Ray. 1963. The Pleistocene vertebrate 
fauna of Reddick, Florida. Quart. Jour. Florida Acad. Sci., vol. 26, 
no. 4, pp. 315-328. 

Hecht, Max K. 1951. Fossil lizards of the West Indian genus Aristelli- 
ger (Gekkcnidae). Amer. Mus. Novitates, no. 1538, pp. 1-33. 



Etheridge: Fossil Lizards 105 

Holman, J. 1958. The Pleistocene herpetofauna of Sabertooth Cave, Citrus 
County, Florida. Copei, no. 4, pp. 276-280. 

. 1959a. A Pleistocene herpetofauna near Orange Lake, Florida. 

Herpetologica, vol. 15, pp. 121-125. 

. 1959b. Amphibians and reptiles from the Pleistocene (Illinoian) 

of Williston, Florida. Copeia, no. 2, pp. 96-102. 

. 1962a. Additional records of Florida Pleistocene amphibians and 

reptiles. Herpetologica, vol. 18, no. 2, pp. 115-119. 

. 1962b. A Texas Pleistocene herpetofauna. Copeia, no. 2, pp. 

96-102. 

. 1963. Late Pleistocene amphibians and reptiles of the Clear Creek 

and Ben Franklin Local Faunas of Texas. Jour. Grad. Res. Cent., vol. 
31, no. 3, pp. 152-157. 

Miller, Gerrit S. 1929. A second collection of mammals from caves near 
St. Michel, Haiti. Smithsonian Misc. Coll., vol. 74, no. 3, pp. 1-8. 

Rao, C. R. 1952. Advanced statistical methods in biometric research. J. 
Wiley and Sons, New York, 390 pp. 

Rouse, Irving. 1964. Prehistory of the West Indies. Science, vol. 144, 
no. 3618, pp. 499-513. 

Schwartz, Albert. 1959. The Cuban lizards of the species Leiocephalus 
carinatus (Gray). Reading Publ. Mus. and Art Gallery Sci. Publ., no. 
10, pp. 1-47. 

Wetmore, Alexander. 1922. Remains of birds from caves in the Republic 
of Haiti. Smithsonian Misc. Coll., vol. 74, no. 4, pp. 1-4. 

Wetmore, Alexander, and Bradshaw H. Swales. 1931. The birds of 
Haiti and the Dominican Republic. Bull. U. S. Nat. Mus., no. 155, 
pp. 234-236. 

Williams, Ernest E. 1960. Notes on Hispaniolan herpetology, 1. Anolis 
cristophei, new species, from the Citadel of King Christophe, Haiti. 
Breviora, no. 117, pp. 1-7. 

Department of Zoology, San Diego State College, San Diego, 
California. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



SODIUM METAPHOSPHATE AND MUTTON TENDERNESS 
D. L. Huffman, A. Z. Palmer, J. W. Carpenter, and R. L. Shirley 

Kamstra (1959) and Carpenter (1961) reported a tenderization 
of fresh ham and beef round by the pre-rigor infusion of sodium 
metaphosphate. Presumably the tenderizing effect was achieved 
by chelating magnesium and calcium ions with the polyphosphate, 
thereby interrupting the normal sequence of rigor. Szent-Gyorgi 
(1951) indicated that magnesium is needed for rigor to proceed 
normally. Several studies have shown actomyosin formation to 
occur during the onset of rigor; other studies indicate that actomyo- 
sin formation is affected by calcium, magnesium, and potassium 
ions as well as adenosine triphosphate. Wierbicki et al. (1954) 
proposed that actomyosin formation might account for the initial 
toughening of meat during rigor. 

Ramsbottom et al. (1949), Paul et al. (1952), Lowe (1955), and 
Wierbicki et al. (1954) showed beef to be tender immediately after 
slaughter but less tender after chilling 24-74 hr. Dodge and Stad- 
elman (1959), working with chickens, found the meat more tender 
when cooked immediately after slaughter than when cooked fol- 
lowing rigor mortis. 

With the establishment of a possible relationship between rigor 
and tenderness it became of interest to determine if the ante- 
mortem injection of chelating materials might alter the normal 
sequence of rigor in such a way as to influence tenderness. The 
purpose of this study was to obtain information on the effects of 
injecting sodium metaphosphate before slaughter on the tender- 
ness of mutton. 

Trial 1 

Procedure. Twenty mature crossbred ewes used in this trial 
were fed a ration consisting of 75 per cent ground snapped corn, 
25 per cent cottonseed meal, and trace mineralized salt at the rate 
of one-half pound per head per day for 1 to 3 weeks before slaugh- 
ter. Coastal Bermuda grass hay was fed, free choice, and water 
was available at all times. 

The arterial occlusion technique suggested by Beuk et al. (1959) 
was practiced on several animals. In this technique, one leg would 
have the circulation tied off and could not receive a tenderizing 



Huffman et al.: Mutton Tenderness 107 

agent injected in the jugular vein. In this manner, it was possible 
to have a control leg and a treated leg from a single animal and 
thus avoid variations in inherent tenderness as well as a differential 
response to treatment by various animals. The sheep were im- 
mobilized by an injection of nembutal in the jugular vein. An 
incision was made, parallel to the L. dorsi, from the anterior edge 
of the hip bone to the 13th rib. The aorta was located and a hemo- 
stat applied immediately posterior to the junction of the common 
iliac and the aorta. The resulting blockage, if properly applied, 
occluded most arterial circulation to the right leg. 

Sodium metaphosphate, (NaP0 3 ) 6 , was combined with distilled 
water to make a solution that contained 10 mg/ml. The pH of 
the solution was approximately 7.4. The trial was divided into 
two phases. The first phase was designed to determine the effect 
of varying levels of sodium metaphosphate on the tenderness of the 
injected leg as compared to the control leg. In phase one 14 ewes 
were divided into seven lots of 2 animals each. Two sheep were 
injected with 1.0 mg sodium metaphosphate/lb. live wt, two 
with 1.5 mg/lb., two with 2.0 mg/lb., two with 2.5 mg/lb., two 
with 3.0 mg/lb., two with 3.5 mg/lb. and two with 4.0 mg/lb. live 
wt. Taste panel tenderness scores of animals in phase one showed 
that 3 mg/lb. live wt of sodium metaphosphate was the more 
effective level. Phase two included 6 animals injected with sodium 
metaphosphate at the rate of 3 mg/lb. live wt. Data collected on 
both phases of this study are combined for presentation in table 1. 

Injections were made with an 18 gauge needle and a glass 
syringe. Animals were slaughtered 5 minutes after the injection 
was completed and the carcasses were chilled for 48 hours at 34- 
36F. All animals subjected to this injection technique were under 
varying degrees of stress prior to slaughter as judged by heart 
rate and respiration rate. 

Two one-inch thick steaks were cut from each leg by cutting 
parallel and immediately posterior to the aitch bone. The steaks 
were broiled in electric ovens to a medium well-done degree. 
Taste panel determinations were made by a two-member panel 
on the top round muscle (semimembranosus). The taste panel 
members rated each sample on a 1 to 9 scale with 1 designating 
too tough to be edible; 2, extremely tough; 3, very tough; 4, below 
average in tenderness; 5, average tenderness; 6, above average — 
tender; 7, very tender — chews easily; 8, extremely tender — grainy; 



108 Quarterly Journal of the Florida Academy of Sciences 





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Huffman et al.: Mutton Tenderness 109 

9, mushy — fibers not distinguishable. This tenderness rating scale 
was used throughout the study. 

Four one-inch thick butterfly chops were removed serially start- 
ing at the posterior end of the loin of each carcass. The chops 
were broiled in electric ovens to a medium well-done (approxi- 
mately 165F.) degree after aging for 48, 72, 96 and 120 hr. The 
chops were evaluated by a two-member taste panel. 

Results. Taste panel evaluations of top round steaks are pre- 
sented in table 1. The first panel member preferred the tenderness 
of sodium metaphosphate injected legs from 12 of the 20 animals, 
the control legs from 2 of 20 animals, and had no preference be- 
tween injected or control legs of 6 animals. The second panel 
member preferred sodium metaphosphate injected legs from 9 of 
17 animals and had no preference on 2 of 17 animals. Averaging 
the preferences, a 54 per cent preference for sodium metaphos- 
phate injected legs was found, whereas there was a 23 per cent 
preference for control legs. No preference was indicated for 23 
per cent of the legs. These data indicate, to a certain extent, that 
the sodium metaphosphate treated legs were more tender than the 
non-treated legs. Tenderness differences noted between the two 
legs were not always in favor of the treated leg which indicates 
a need for further investigation of the suggested tenderizing effect. 

Panel tenderness evaluations on loin chops are shown in table 
1, These chops were taken from an area of the carcass subject 
to effects of the ante-mortem injection treatments. The purpose 
of testing the chops as to tenderness was to train the panelists in 
testing mutton chops for tenderness and to determine the level of 
metaphosphate giving the most promising tenderness response. 
On the basis of these data it appeared that the 3 mg sodium meta- 
phosphate/lb. live wt was the most desirable level to use in fur- 
ther studies. 

Trial 2 

Procedure. Thirty-two mature crossbred ewes were used in 
this trial. The ewes, obtained at the same time as those in trial 1, 
were fed and cared for in the same manner as the animals used in 
trial 1. 

Trial 2 was designed to provide data that would either verify 
or reject treatment effects noted in trial 1. Data obtained from 
the 8 animals injected with 3 mg sodium metaphosphate/lb. live 
wt in trial 1 are included in this trial also. 



110 Quarterly Journal of the Florida Academy of Sciences 

Four lots with 8 animals per lot were used in this trial. Animals 
were allotted by age and weight. The animals in lot 1 were not 
injected and served as controls. Lot 2 was made up of 8 animals 
from trial 1 that were injected with sodium metaphosphate, 3 
mg/lb. live wt, following nembutal injection and arterial occlu- 
sion. The tenderness data on the leg not subjected to the phos- 
phate treatment by virtue of its arterial occlusion are not included 
in trial 2. The reason for including data from these 8 animals 
was to find out by a more direct comparison (trial 2, lot 2 vs. lot 
4) if the observed tenderization were due to metaphosphate or 
other factors incidental to arterial occlusion. The third group of 
animals received sodium metaphosphate in the same total amout 
(3 mg/lb. live wt) as other injected lots; however, the solution was 
divided into three equal doses and administered 125, 65, and 5 
minutes ante-mortem. The animals in lot 4 received a single in- 
jection of sodium metaphosphate, at the rate of 3 mg/lb. live wt, 
administered 5 minutes ante-mortem. It should be noted that the 
difference between lots 2 and 4 lies in the fact that the sheep in 
lot 2 underwent nembutal injection and surgical blockage of the 
right leg prior to the sodium metaphosphate injection, while lot 
4 animals received only the metaphosphate injection. 

Sodium metaphosphate solution was prepared in manner de- 
scribed for trial 1. It should be noted that solutions were dis- 
carded after each use and new solutions prepared to avoid possible 
bacterial contamination. 

Injections were made as previously described. Sheep were 
slaughtered and the carcasses were chilled at 34-36F. for 48 hours. 

Leg steaks and loin chops were removed in the same manner 
and at the same time intervals after slaughter as indicated for trial 
1. The taste panel members were the same as those used in trial 1. 
Tenderness rating was on the 1 to 9 scale described for trial 1. 

Trial 2 tenderness data on the top round steak were statisti- 
cally treated by analysis of variance as described by Snedecor 
(1956). Trial 2 tenderness data on the loin chops were statistically 
analyzed by analysis of variance according to Henderson (1959). 

Results. The results of the taste panel evaluation of top round 
steaks are shown in table 2. The treatment means appear to be 
sufficiently different to be significant; however, such was not the 
case. Despite the lack of statistical significance it should be noted 
that only 2 out of 24 animals in lots 2, 3, and 4 were below aver- 



Huffman et al.: Mutton Tenderness 111 

age in tenderness, while 5 of the 8 animals in the non-injected 
control lot were below average in tenderness. A rather close agree- 
ment may be noted between the tenderness values for top round 
steaks and loin chops. 

The results of the taste panel evaluations of the loin chops are 
shown in table 2. A significant difference (P<0.01) in the tender- 
ness of chops aged for different periods of time was obtained. 
Differences in tenderness between treatments were not significant. 
It is generally accepted that aging allows tenderization; it would 
have been interesting, though, had there been a significant inter- 
action between injection treatment and the aging effect on ten- 
derness. 

The most strongly suggested tenderizing effect from treatment 
was in lot 2; these animals had received nembutal and were under 
considerable stress during the arterial occlusion operation. The 
tenderizing effect, seen in average tenderness values but lacking 
statistical significance, may have been due, in part, to the nembu- 
tal or to the physiological shock. Such interpretation seems valid 
in view of the fact that while lot 2 was similar to lot 4 in ante- 
mortem injection level, lot 4 chops were not improved in tender- 
ness over the control chops as was lot 2. 

Lot 3 and 4 animals received the same amounts of sodium 
metaphosphate; lot 3 animals were treated in 3 equal doses at 125, 
65, and 5 minutes ante-mortem, where lot 4 animals were treated 
in one injection 5 minutes ante-mortem. The treatment for lot 3 
was more effective in tenderizing than the treatment for lot 4 as 
seen in average values presented in table 2. For this reason, a 
longer interval of time between injection and slaughter appears 
to be a logical modification in method for future studies. 

Summary 

Two trials were conducted to develop experimental techniques 
and gain knowledge concerning the usefulness of sodium meta- 
phosphate as a tenderizing agent when injected ante-mortem. 

The first preliminary trial with mature sheep provided an 
opportunity to study an arterial occlusion technique which made 
it possible to study effects of ante-mortem injections on a "within 
animal" basis, thereby holding constant inherent tenderness differ- 
ences that are known to exist between animals. The arterial oc- 



112 Quarterly Journal of the Florida Academy of Sciences 



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Huffman et al.: Mutton Tenderness 113 

elusion technique made it possible to gain more information from 
a smaller number of animals than would have been required other- 
wise. In trial 1 it was learned that loin chops from sheep were 
more uniform in tenderness with the techniques of cooking and 
sampling used in this study and thereby permitted a more valid 
estimate of the tenderness of an animal than leg steaks. Loin chops 
from animals injected at the rate of 3 mg/lb. live wt appeared to 
be more tender than chops from animals injected at lower or 
higher levels. There was a decided tendency for the leg steak 
receiving sodium metaphosphate by ante-mortem injection to be 
more tender than the corresponding control leg steak that was not 
subjected to the sodium metaphosphate. 

In trial 2, the ante-mortem injection of sodium metaphosphate 
did not effect the tenderness of leg steaks to the extent of statisti- 
cal significance. It is noted, however, that leg steaks from animals 
subject to injection were on an average more tender than steaks 
from control animals. The ante-mortem treatment of this study 
did not significantly effect loin chop tenderness. On an aver- 
age basis, however, loin chops from animals injected with so- 
dium metaphosphate following the arterial occlusion operation 
and slaughtered 5 minutes later were more tender than chops from 
control animals. Further, chops from animals injected with sodium 
metaphosphate 125, 65, and 5 minutes ante-mortem were more 
tender than chops from control animals on the average. 

Acknowledgment 

This study was supported in part by donation of the experi- 
mental animals by Lykes Packing Company, Tampa, Florida. 

Literature Cited 

Beuk, J. F., A. L. Savich, and P. A. Goeser. 1959. Method of tendering 
meat. United States Patent 2,903,362. 

Carpenter, J. A., R. L. Saffle, and L. D. Kamstra. 1961. Tenderization 
of beef by pre-rigor infusion of a chelating agent. Food Technol., vol. 
15, pp. 197-198. 

Dodge, J. W., and W. J. Stadelman. 1959. Post mortem aging of poultry 
meat and its effect on the tenderness of the breast muscles. Food 
Technol., vol. 13, pp. 81-84. 



114 Quarterly Journal of the Florida Academy of Sciences 

Henderson, C. R. 1959. Design and analysis of animal husbandry experi- 
ments. Techniques and Procedures in Animal Production Research, 
Amer. Soc. Animal Production Monograph, pp. 2-55. 

Kamstra, L. D., and R. L. Saffle. 1959. The effects of a pre-rigor in- 
fusion of sodium hexametaphosphate on tenderness and certain chemi- 
cal characteristics of meat. Food Technol., vol. 13, pp. 652-655. 

Lowe, B. 1955. Experimental Cookery. John Wiley and Sons, Inc., New 
York. 

Paul, P., L. J. Bratzler, E. D. Farwell, and K. Knight. 1952. Studies 
on tenderness of beef, I. Rate of heat penetration. Food Res., vol. 
17, pp. 504-510. 

Ramsbottom, J. M., and E. J. Strandine. 1949. Initial physical and chem- 
ical changes in beef as related to tenderness. Jour. Animal Sci., vol. 8, 
pp. 398-410. 

Snedecor, G. W. 1956. Statistical methods. Iowa State College Press, 
Ames, ed. 5, 534 pp. 

Szent-Gyorgi, A. 1951. Chemistry of muscular contraction. Academic 
Press, Inc., New York. 

Wierbicki, E., L. E. Kunkle, V. R. Cahill, and F. E. Deatherage. 1954. 
The relation of tenderness to protein alterations during post-mortem 
aging. Food Technol., vol. 8, pp. 506-511. 

Department of Animal Science, University of Florida, Gaines- 
ville, Florida. Florida Agricultural Experiment Stations Journal 
Series No. 2084. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



BIRD REMAINS FROM A KENTUCKY INDIAN MIDDEN 
Glen E. Woolfenden 

In 1939 bones and artifacts were removed from an Indian mid- 
den discovered near Perkins Creek on the Will Rottering farm at 
Paducah, McCracken County, Kentucky, by Dr. Andrew L. Pick- 
ens of Paducah Junior College and Mr. Fain King from Wickliffe 
(Anonymous, 1939). The midden lay in the path of the proposed 
Section A of the city noodwall (Scott, 1939), and the material was 
gathered rapidly and stored in the Junior College. In 1958 and 
1961 with the cooperation of Royce H. Gregory, the business man- 
ager, and Dr. R. G. Matheson, the president, I removed all the 
bird bones and some mammal bones and artifacts for study; the 
remaining material was discarded. 

The collection contains specimens of 22 species of birds. Spe- 
cific identification was based in part on the geographic ranges of 
living species. The list includes the pied-billed grebe (Podilymbus 
podiceps), Canada goose (Branta canadensis), mallard (Anas platy- 
rhynchos), green- winged teal (Anas carolinensis), blue-winged teal 
(Anas discors), canvasback (Ay thy a valisineria), lesser scaup (Ay thy a 
affinis), red-tailed hawk (Buteo jamaicensis), greater prairie chicken 
(Tympanuchus cupido), bobwhite (Colinus virginianus), turkey 
(Meleagris gallopavo), American coot (Fulica americana), passenger 
pigeon (Ectopistes migratorius), screech owl (Otus asio), great 
horned owl (Bubo virginianus), barred owl (Strix varia), yellow- 
shafted flicker (Colaptes auratus), pileated woodpecker (Dryocopus 
pileatus), red-headed woodpecker (Melanerpes erythrocephalus), 
common raven (Corvus corax), common crow (Corvus brachyrhyn- 
chos), and the family Icteridae, probably the common grackle 
(Quiscalus quiscida). 

Of special interest is the occurrence of the partial right humerus 
of the common raven. According to Mengel (in press) the species 
was probably once resident through much of the Cumberland 
Plateau and Mountains, and appeared elsewhere in Kentucky as 
a vagrant. However, the species disappeared from the state dec- 
ades ago and no specimen is extant. The bone is larger in shaft 
width and depth than eleven specimens of Corvus corax sinuatus 
in the University of Kansas Museum of Natural History and is 
closer to the one specimen of C. c. principalis, which is the race to 



116 Quarterly Journal of the Florida Academy of Sciences 

be expected in Kentucky. The remaining 21 species are all known 
to occur or have occurred in western Kentucky, although specimens 
of several, the greater prairie chicken for example, apparently do 
not exist (Mengel, op. cit.). 

Various artifacts from the site were examined by Dr. Berle 
Clay, Museum of Anthropology, University of Kentucky, who esti- 
mated they came from the middle or latter half of the Mississippian 
Period somewhere between 1300 and 1500 A.D. (letter March 13, 
1964). Avian material from the Wickliffe Mounds, located in Bal- 
lard County, and of approximately the same age as the Paducah 
midden was examined by Dr. Paul W. Parmalee (letter March 30, 
1964), who identified bones from eight species of birds including 
the following forms not found at the Paducah site: American 
widgeon (Mareca americana), wood duck (Aix sponsa), bald eagle 
(Haliaeetus leucocephalus), and sandhill crane (Grus canadensis). 
The turkey was by far the most abundant bird in the Paducah ma- 
terial. At Wickliffe, however, the species was outnumbered by 
bones of the mallard or black duck (Anas rubripes). 

Literature Cited 

Anonymous. 1939. Volunteer workers sought to save valuable Indian 
mound. The Paducah Sun-Democrat, September 17, 1939. 

Mengel, Robert M. In press. The birds of Kentucky. Ornith. Monogr. no. 
2, Amer. Ornith. Union. 

Scott, Burgess. 1939. Off the record. The Paducah Sun-Democrat, Sep- 
tember 18, 1939. 

Department of Zoology, University of South Florida, Tampa, 
Florida. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



A NEW GECKO FROM THE VIRGIN ISLANDS 

Richard Thomas 

During a collecting trip to the Virgin Islands in the summer of 
1964 the xeric but wooded hillslopes of Virgin Gorda yielded not 
only the ubiquitous Sphaerodactylus macrolepis Gunther but an- 
other, undescribed diminutive species, here named 



Sphaerodactylus parthenopion, new species 



Type. MCZ 77211 (original number ASFS V3734), an adult 
female, collected on hillside above Pond Ray, Virgin Gorda, Rrit- 
ish Virgin Islands, on 12 August 1964, by Richard Thomas (Fig. 1). 





Fig. 1. Dorsal body and head pattern of the type specimen of Sphaero- 
dactylus parthenopion (MCZ 77211). 

Fig. 2. Variant head pattern of Sphaerodactylus parthenopion (ASFS 
X3658); one specimen has both ends of cephalic bar connected to postocular 
stripes. 

Fig. 3. Dorsal body and head pattern typical of many Sphaerodactylus 
nicholsi (ASFS X4246). 

Paratypes. ASFS V3658-59, ca. 0.5 mi. N Pond Ray, 10 August 
1964, R. Thomas; ASFS V3664, Pond Ray, 10 August 1964, R. Thom- 
as; ASFS V3681, between Little Dix Ray and Savana Ray, 11 Au- 
gust 1964, R. Thomas; AMNH 92821, ca. 0.5 mi. N Pond Ray, 11 
August 1964, R. Thomas; AMNH 92822-24, hillside above Pond 



118 Quarterly Journal of the Florida Academy of Sciences 

Bay, 12 August 1964, R. Thomas; MCZ 77212-14, ca. 0.5 mi. N 
Pond Bay, 13 August 1964, R. Thomas; KU 79852-53, between Lit- 
tle Dix Bay and Savana Bay, 15 August 1964, David C. Leber and 
R. Thomas; KU 79854, SW slope of Gorda Peak, ca. 500', 16 Au- 
gust 1964, R. Thomas. All above localities are on the island of 
Virgin Gorda. 

Diagnosis. A species of Sphaerodactylus characterized by its 
very small size; small, but keeled and imbricate dorsal scales; lack 
of a middorsal zone of granules or granular scales; a generally 
uniform and dark dorsal coloration; lack of scapular or sacral pat- 
tern; and a distinctive cephalic pattern as described below. 

Distribution. Known only from the island of Virgin Gorda. 

Description of type. Dorsal scales small, acute to rounded, 
moderately keeled and slightly imbricate; 32 scales from axilla to 
groin counted dorsolaterally. Some crowding and reduction in 
size of scales middorsally but no zone of granules or granular 
scales present. Granular scales of top of head and anterior neck 
becoming flattened and imbricate at mid-neck. Gular and pectoral 
scales keeled. Ventral scales obtuse to rounded, smooth and im- 
bricate; 28 scales from axilla to groin along mid ventral line. Scales 
around midbody 52. Internasals 2; upper labials to middle of eye 
3. Lamellae on fourth toe of right foot 8. Dorsal scales of tail 
acute, keeled, and erect from tail; tail regenerated. Snout- vent 
length 17 mm. 

Dorsal ground color of head and body deep brown. A narrow, 
dark-edged, yellow-brown postocular stripe extends over each tem- 
ple and fades out at base of head. A faint, preocular light bar 
across base of snout. A dark-edged, more or less oval, transverse 
yellow-brown bar present on top of head just behind eyes, its ends 
not reaching laterally to postocular stripes. A faint, light occipital 
spot present. Dorsal body coloration of scattered dark brown or 
black scales with a slight concentration of dark scales along mid- 
dorsal line. Dorsa of hindlimbs with irregular mottling of darker 
scales. Ground color of tail yellowish brown, pattern of irregular 
short linear or clustered dark elements. Ventral ground color 
light, grayish to cream. Gular pattern a faint reticulum of dark 
pigmentation; lateral light areas of gular pattern continuous with 
light markings on upper labials and temporal region which radiate 
from eye. Brown dorsal coloration invades venter, fading out 
centrally except for dark edges to many scales. 



Thomas: Virgin Island Lizard 119 

Variation. Scalation of the 14 paratypes is much the same as 
that of the type. Dorsal scales from axilla to groin range from 
30-35 in 12 specimens counted. Ventral scales axilla to groin 26- 
31; scales around midbody 50-55. Throat and pectoral scales are 
keeled in all but one specimen. Fourth toe lamellae 8 (mode) or 
9. Upper labials typically 3 to mid-eye; 4 specimens have only 2 
upper labials on one side. In specimens with unregenerated tails 
the tips of the scales stand out somewhat from the surface of the 
tail imparting a roughened appearance. Escutcheons of the five 
adult males vary in length from 3-5 scales and in width from 11- 
13 scales. Snout-vent lengths vary from 12-18 mm (length of the 
three largest females). 

The coloration of the paratypes is much the same as that of 
the type. The overall impression of the color of these lizards is 
that they are a dark, velvety brown. Some specimens have a uni- 
form reticulum of darker scales instead of the irregularly scattered 
dark scales of the type (the modal coloration); others are a nearly 
uniform brown with few or no darker scales. The cephalic pat- 
terns are also generally much like that of the type; two specimens 
have the cephalic bar joined at one or both ends with a postocular 
stripe (Fig. 2). One of these specimens has diffuse, light transverse 
markings on the neck. The preocular transverse bar is evident in 
all specimens, albeit faintly is some. The pattern of light lines 
which radiate from the eye to the underside of the head is more 
prominent than in the type in half of the paratypes, and as prom- 
inent or less so in half. The gular pattern is very faint in some 
specimens, but in others it is composed of rather prominent linear 
elements which converge towards the midline from either side and 
continue posteriorly, ending at the neck. 

Comparisons. The most pertinent comparison for the new 
form is with Sphaerodactylus nicholsi Grant, a species presently 
known only from the southwestern corner of Puerto Rico (Grant, 
1931; Albert Schwartz, field data). S. nicholsi is similar to parth- 
enopion in its small size and related color pattern and is doubtless 
its closest relative. S. nicholsi possesses a crescentic, light-colored 
cephalic pattern (Fig. 3) which may meet the postocular stripes 
laterally; this pattern is not present in all specimens. The post- 
ocular stripes of nicholsi continue onto the body, sacrum and tail 
with a few exceptions when stripes of any kind are virtually ab- 
sent. Other than occasional faint dorsolateral stripes on the prox- 



120 Quarterly Journal of the Florida Academy of Sciences 

imal part of the tail, no specimens of parthenopion possess stripes 
beyond the postocular stripes ending on the neck. About 60 per 
cent of the specimens of nicholsi possess a scapular pattern of two 
small, light spots surrounded by a zone of black; nicholsi also has 
a dark-edged U- or Y-shaped sacral pattern formed by the conflu- 
ence of the dorsolateral stripes. S. parthenopion possesses neither 
of these pattern elements. S. nicholsi attains a larger size than 
parthenopion; aside from the fact that large specimens attain snout- 
vent lengths of 20-22 mm (versus a maximum of 18 mm for par- 
thenopion), nicholsi is a bulkier lizard in general appearance. 
S. parthenopion is certainly the smallest known Splwerodactylus, 
and it must rank among the smallest of lizards. 

Scalation differences between the two forms are striking and 
reflect the smaller size of dorsal scales in parthenopion, a. fact 
which is evident from mere inspection. Table 1, comparing body 
scale counts of 12 parthenopion with counts taken on 22 specimens 
of nicholsi, suggests the differences in scale size. 

TABLE 1 
Scale counts of Sphaerodactylus 



Body scales 



S. nicholsi 



S. parthenopion 



Dorsal scales axilla to groin 
Ventral scales axilla to groin 
Scales around midbody 



19-24 
21-26 
35-42 



30-35 
26-29 
50-55 



Of the 15 specimens of S. parthenopion only one specimen has but 
a single internasal, the rest having either 2 (mode) or 3. On the 
other hand, of 41 specimens of S. nicholsi, all have a single inter- 
nasal but 4 which have 2 internasals. Escutcheons of male nicholsi 
attain larger size than do those of parthenopion, but there is con- 
siderable overlap. 

Sphaerodactylus parthenopion may be told from S. macrolepis 
with which it occurs syntopically by the much larger size of the 
latter (adults 25-30 mm snout-vent) and by the latter's larger and 
coarser scales. S. macrolepis has a pattern of dark lateral stripes 
and dorsal spotting on a tan or light brown ground color with a 
boldly black-edged pair of scapular spots (females) or a nearly uni- 
form yellow-brown body color, weak or absent scapular pattern, 



Thomas: Virgin Island Lizard 121 

and contrasting head pattern of black vermiculations on a gray- 
ground color or unicolor yellow or orange heads (males). 

The new species was not obtained on any of the other Virgin 
Islands visited (St. Croix, St. Thomas, St. John, Tortola, Anegada, 
and assorted smaller islets) nor during a brief visit to Vieques and 
Culebra. Thus it appears that Sphaerodactylus parthenopion is 
separated from its nearest relative by the intervening majority of 
the Virgin Islands plus Vieques and Culebra (Virgin Gorda is 
among the easternmost of the Virgin Islands). It is of interest to 
note that the toad Bufo lemur Cope is known only from Puerto 
Rico and Virgin Gorda, though in this case differentiation to even 
subspecific level has apparently not taken place (Ruibal, 1959, 
p. 10). 

S. parthenopion was collected primarily on the very rocky hill- 
sides of the hilly portions of the island. These hillsides are cov- 
ered with a low, xeric woods interspersed with cacti and thorn 
scrub. The geckos were not observed to "swarm" in the leaf litter 
as did the sympatric S. macrolepis; they were found only by turn- 
ing rocks, and then not very commonly. S. parthenopion was taken 
once at a sea level locality (Pond Bay), but in its characteristic rocky 
habitat, and was not observed in the beach area among the sea- 
grape litter. On the lower-lying southern portion of the island 
where S. macrolepis abounded and where large series were taken 
in piles of rotting palm trash, the smaller species was not found. 
The Puerto Rican relative, S. nicholsi, apparently does not occupy 
the same or so restricted a habitat as parthenopion, for it can also 
be found abundantly in the leafy floor of coastal xeric woods or 
beach growth (Grant, 1931; A. Schwartz, field notes). 

I wish to thank Dr. Albert Schwartz, who sponsored the field 
work in the Virgin Islands and the study of these geckos, and Mr. 
David C. Leber for his assistance in the field. 

Specimens Examined 

The following symbols are used to designate collections in 
which the type series of Sphaerodactylus parthenopion is housed: 
AMNH (American Museum of Natural History), ASFS (Albert 
Schwartz Field Series), KU (University of Kansas Museum of Nat- 
ural History), MCZ (Museum of Comparative Zoology, Harvard). 

Sphaerodactylus parthenopion: As listed for type and para- 
types. 



122 Quarterly Journal of the Florida Academy of Sciences 

Sphaerodactylus nicholsi: Puerto Rico: ASFS X4199-205, 8 
km SE Guanica; ASFS X4206-12, Playa de Cana Gorda; ASFS 
X4213-27, 3 mi. SW Ensenada; ASFS X4250, 3 mi. SW Ensenada; 
ASFS X4280-84, 9 km SE Guanica; ASFS X4286-87, 10 km SE 
Guanica; ASFS X4362, Laguna Cartagena, west end. 

ASFS specimens of S. townsendi Grant, S. gaigeae Grant, S. 
macrolepis Gunther, and S. m. parvus King were casually examined 
in the initial search for species possibly related to S. parthenopion. 

Literature Cited 

Grant, Chapman. 1931. The sphaerodactyls of Porto Rico, Culebra and 
Mona islands. Jour. Dept. Agri. Puerto Rico, vol. 15, no. 3, pp. 
199-213. 

Ruibal, Rodolfo. 1959. Bufo gundlachi, a new species of Cuban toad. 
Breviora, no. 105, pp. 1-14. 

10000 SW 84th Street, Miami, Florida. 



Quart, Jour. Florida Acad. Sci. 28(1) 1965 



NEREID BLISTERS IN FLORIDA SCALLOPS 

Harry W. Wells 

The nereid polychaete Ceratonereis tridentata (Webster) has 
been implicated as a pest of calico scallops, Aequipecten gibbus 
(Linne), in North Carolina waters (Wells and Wells, 1962). This 
polychaete was found in large, dark-colored blisters which the 
scallop had been stimulated to secrete on the inner surfaces of its 
valves. In the present report, Ceratonereis tridentata is recorded 
from the calico scallop in Florida waters, and data on its incidence 
are presented. 

Materials and Methods 

Through the courtesy of Capt. J. S. Andrew and Mr. John Salva- 
dor of S. Salvador Sons Seafood Market, St. Augustine, Florida, 
one bushel of calico scallops was obtained from a commercial 
trawler for examination of the associated organisms. These scal- 
lops had been dredged January 20-25, 1964, from depths of 15-25 
fathoms in the scallop beds east of Cape Canaveral (Cape Ken- 
nedy), Florida. The scallops were put on ice on board the trawler, 
washed in fresh water at dockside, and returned to ice for trans- 
port to the laboratory. Each scallop was shucked and examined 
for the presence of blisters caused by C. tridentata. For each 
blister, a careful examination was made under a stereoscopic mi- 
croscope in order to locate and identify the inhabitant. The ex- 
amined scallops ranged from 50-61 mm in length, with a mean 
of 56.2 mm. 

Supplementary information on the spatial distribution of Cera- 
tonereis blisters was obtained by analyzing the location of blisters 
in a dozen additional shell pairs. These shells, which were se- 
lected from the discarded shells accumulated outside the shucking 
plant, had been dredged during the preceding two months (De- 
cember, 1963, and January, 1964) from the same beds off Cape 
Canaveral. 

The Cape Canaveral scallop beds encompass an extensive off- 
shore area extending from approximately the latitude of Ormond 
Beach southward to the latitude of Stuart (Bullis and Cummins, 
1961). The beds are generally between 15-30 fathoms depth in 
this area, where the bottom is composed principally of sand and 
shell. Water temperatures from 19.7-28. 8C and salinities from 33.5 



124 Quarterly Journal of the Florida Academy of Sciences 

to 36.3 o/oo have been recorded for this area (Anderson, et al., 
1961a, 1961b). 

Observations 

Of 418 calico scallops examined, blisters that could be attrib- 
uted to Ceratonereis tridentata were found in 14 scallops (3.3 per 
cent). Specimens of C. tridentata up to 65 mm long were removed 
from several blisters. In several other cases where no worm was 
found, the blister inhabitant had recently vacated the blisters, 
probably during handling and transportation prior to examination. 
Superficially similar blisters, which were found in six scallops (1.4 
per cent) had been caused by the spionid polychaete Polydora 
websteri. In addition, most shells had small blemishes on the in- 
terior surface caused by incomplete perforations of the shell by 
P. websteri. 

The localization of the Ceratonereis blisters contained in these 
scallop shells is analyzed in Table 1, with supplementary informa- 
tion on the position of 12 blisters recovered from the shell ac- 
cumulations at the shucking plant. Most blisters were located in 

TABLE 1 

Location of Ceratonereis blisters in Aequipecten gibbus from off 
Cape Canaveral. Florida 











Per 


cent 




Upper 


Lower 






North 


Position 


valve 


valve 


Totals 


Florida 


Carolina * 


Umbonal 


20 


4 


24 


92 


65 


Marginal 





2 


2 


8 


35 


Anterior 
opening 


19 


5 


24 


92 


81 


Posterior 
opening 





1 


1 


4 


1 


Anterior & 












posterior 


1 





1 


4 


18 


openings 












Totals 


20 


6 


26 


100 


100 


Per cent 


77 


23 


100 






N.C. (%) 


75 


25 


100 







North Carolina data from Wells and Wells (1962). 



Wells: Blisters in Scallops 125 

an umbonal position in the upper shell, i.e. occupying the concavity 
under the umbo, dorsal to the adductor muscle. These blisters 
typically possessed an opening to the exterior near the byssal notch, 
where the anterior wing of the scallop shell joins the anterior 
shell border. 

Typical blisters were approximately 30 mm long by 20 mm 
high, ovoid to irregular in shape, and elevated 2-4 mm above the 
normal shell surface. Their walls were formed of a thin layer 
of dark brown conchiolin, which in some cases had been obscured 
partially by subsequent deposits of nacreous shell material. In 
several cases, a neat circular perforation of the blister wall pro- 
vided direct access to the scallop's mantle cavity; in other cases, 
such an opening may have been overlooked or destroyed in shuck- 
ing. Coarse detrital material contained within the blister often 
contributed an even darker quality to its appearance. The detrital 
material contained within blisters caused by C. tridentata is usu- 
ally distinctly coarser than that which is contained within blisters 
caused by P. websteri. Frequently, a flimsy mucous-like tube 2-3 
mm wide occupied much of the space within the blister. When 
a specimen of C. tridentata was located, it was contained within 
such a tube. 

Discussion 

Hartman (1945, 1951) recorded Ceratonereis tridentata as oc- 
curring in shelly bottoms from New Jersey to Texas, and provided 
adequate descriptions of the species. However, no indication of 
its role as a pest of pelecypods had appeared in scientific litera- 
ture. Its recognition as a pest of Aequipecten gihbus resulted from 
analysis of its abundance and accompanying morphological anom- 
alies in scallops from off the North Carolina coast (Wells and Wells, 
1962). Because it can harm its host, C. tridentata may be regarded 
as a parasite of the calico scallop, although most biologists may 
prefer to call it a facultative commensal. In the North Carolina 
study, Ceratonereis blisters occurred in a much larger proportion 
of the scallop population (24 versus 3.3 per cent for this Florida 
population). On the basis of its incidence in this collection, C. 
tridentata would not be regarded as a serious pest on the east 
coast of Florida. 

For the location of Ceratonereis blisters in the scallop shell, 
comparison with data from the North Carolina collection is pro- 



126 Quarterly Journal of the Florida Academy of Sciences 

vided in Table 1. There is very close agreement for the proportion 
of blisters in upper valves as opposed to lower valves, and good 
agreement on the position of exterior openings. The most signifi- 
cant difference in Ceratonereis blisters was the low incidence of 
marginal blisters in the Florida scallops, much lower than in the 
North Carolina collection. Coincidentally, the incidence of Poly- 
dora blisters in these Florida scallops was very low, whereas 
Fohjdora blisters occurred in 99 per cent of the North Carolina 
shells and usually occurred in a marginal position. Thus, the 
principal differences in the Florida and North Carolina collections 
were (1) the lower overall incidence of Ceratonereis blisters, (2) 
the lower proportion of marginal blisters, and (3) the much lower 
incidence of Polydora blisters. 

Only circumstantial evidence is available for relating these 
differences to environmental factors; experimental analysis of the 
various relationships is lacking. The lower incidence of Ceratone- 
reis blisters and Polydora blisters might be related to differences 
in size or condition of the scallops, in size, condition, or abundance 
of the invading polychaetes, or in physical differences in the en- 
vironment. In the North Carolina study, where marginal blisters 
occurred in 35 per cent of the affected scallops, several scallops 
were found in which sand and debris had been forced between 
the mantle and shell in the course of dredging operations. Intro- 
duction of such foreign material under the mantle would make 
the scallop more susceptible to exploitation by these invasive poly- 
chaetes. Consequently, fishing pressure on the scallop population 
has been cited as a factor favoring or improving conditions for the 
production of Ceratonereis blisters. Moreover, the conditions pro- 
duced by dredging operations would particularly favor the pro- 
duction of marginal blisters (as opposed to umbonal blisters) as a 
reaction to either polychaete species. Because the North Carolina 
scallop population had been exposed to considerable fishing pres- 
sure from commercial fishing trawlers during the preceding year, 
while the Florida population had not, much of the difference in 
blister incidence may be attributable to these differences in fish- 
ing pressure. 

The relationship between Ceratonereis tridentata and the calico 
scallop has been described and discussed by Wells and Wells 
(1962). The blisters associated with C. tridentata are formed by 
the scallop's mantle in response to its irritation by this polychaete. 



Wells: Blisters in Scallops 127 

After the worm's invasion between shell and mantle, deposits of 
conchiolin more or less isolate the invader from the scallop tissues. 
In later stages, overlying nacreous shell depoits reinforce the blister 
wall. Not only does blister construction distort the scallop's sym- 
metry, but the essential deposition of conchiolin and of extra shell 
material competes with normal growth and reproduction for the 
scallop's energy. In many cases, the gills of an infested scallop 
are eroded so that feeding is impaired, and extreme cases of gill 
erosion may be accompanied by atrophy of the gonadal mass. In 
this manner, C. tridentata reduces its host's reproductive poten- 
tial and weakens its host, perhaps making it less able to resist 
other enemies. 

Sequestered in its blister, C. tridentata certainly receives pro- 
tection from potential predators to which it would be exposed on 
the outer shell surfaces. Free-living specimens have been found 
on the outside shell surfaces of the calico scallop, almost hidden 
in protecting crevices among the epifauna (Wells, et al., 1964). 
Those specimens of C. tridentata which have access to the host's 
mantle cavity through a perforation of the blister wall probably 
obtain food in the mantle cavity. It has been suggested (Wells 
and Wells, 1962) that they might steal food from the scallop by 
intercepting the strings of mucous and food filtered by the gills as 
these materials are conveyed toward the mouth. Occupants of 
blisters with no direct access to the mantle cavity presumably ob- 
tain their food from another source. The scallop epifauna, which 
provides a degree of protection for some individuals, may also 
provide their food. 

Examination of similar blisters in the bay scallop Aequipecten 
irradians from inshore Florida waters has not revealed specimens 
of C. tridentata. As in the calico scallop, large, typical Polydora 
blisters may occur in the bay scallop, and they may secondarily be 
invaded by other nereid polychaetes which superficially resemble 
C. tridentata. Consequently, careful examination and identifica- 
tion of the blister inhabitant is necessary for distinguishing be- 
tween blisters caused by C. tridentata and those caused by other 
agents. 

The effect of C. tridentata on the calico scallop fishery in the 
vicinity of Cape Canaveral must be regarded as being very small, 
because (1) the incidence of Ceratonereis blisters is low, (2) there 
is no evidence that this species removes scallops from the popu- 



128 Quarterly Journal of the Florida Academy of Sciences 

lation, and (3) the scallop fishery itself is small and erratic. Never- 
theless, recognition of this species is important because (1) its 
effects may increase with greater fishing pressure, (2) it may reduce 
the reproductive potential of the scallop population, and (3) by 
weakening scallops, its invasion may aid other enemies of the 
calico scallop. 

Acknowledgments 

This study was made in conjunction with a more comprehen- 
sive study of sponges of the Carolinian Province, supported by 
National Science Foundation Grant BG-128 and the Florida State 
University Research Council. The author gratefully acknowledges 
the assistance of Mrs. Mary Jane Wells in all phases of this study. 

Literature Cited 

Anderson, W. W., J. E. Moore, and H. R. Gordy. 1961a. Water tempera- 
tures off the south Atlantic coast of the United States, THEODORE 
N. GILL cruises 1-9, 1953-54. Fish & Wildl. Serv. Spec. Sci. Rept. 
Fish., no. 380, pp. 1-206. 

. 1961b. Oceanic salinities off the south Atlantic coast of the United 

States, Theodore N. Gill cruises 1-9, 1953-54. Fish & Wildl. Serv. 
Spec. Sci. Rept. Fish., no. 389, pp. 1-207. 

Bullis, H. R., and R. Cummins, Jr. 1961. An interim report on the Cape 
Canaveral scallop bed. Commerc. Fish. Rev., vol. 23, no. 10, pp. 1-8. 

Hartman, Olga. 1945. The marine annelids of North Carolina. Duke 
Univ. Mar. Sta. Bull., no. 2, pp. 1-51. 

. 1951. The littoral marine annelids of the Gulf of Mexico. Publ. 

Inst. Mar. Sci. Texas, vol. 2, no. 1, pp. 7-124. 

Wells, H. W., and M. J. Wells. 1962. The polychaete Ceratonereis tri- 
dentata as a pest of the scallop Aequipecten gibbus. Biol. Bull., vol. 
122, pp. 149-159. 

Wells, H. W., M. J. Wells, and I. E. Gray. 1964. The calico scallop 
community in North Carolina. Bull. Mar. Sci. Gulf & Carib., vol. 14, 
no. 4, pp. 561-593. 

Department of Biological Sciences, Florida State University, 
Tallahassee, Florida. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



MOLYBDENUM TOXICITY IN RATS AND RABBITS 
L. R. Arrington, C. B. Ammerman, and J. E. Moore 

One of the characteristic symptoms of molybdenum toxicity in 
animals is retarded growth or poor weight gain. This effect and 
other symptoms of molybdenum toxicity have been demonstrated 
in several species and the reports have been reviewed by Dick 
(1956), Underwood (1962), and Miller and Engel (1963). Data are 
limited, however, which relate the poor growth to feed consump- 
tion and feed utilization. Johnson and Miller (1961) have reported 
that 600 ppm dietary molybdenum, when fed to rats in a pair- 
feeding trial, resulted in reduced feed consumption and efficiency 
of feed utilization. In a subsequent study, these authors (1963) 
demonstrated that rats fed molybdenum utilized nutrients less effi- 
ciently, but digestion and absorption of nitrogen were not affected. 
They proposed the hypothesis that molybdenum may inhibit some 
facet of protein synthesis. Monty and Click (1961) have demon- 
strated that rats develop a sensory recognition of molybdenum in 
the diet and reject diets containing toxic levels of the element. 
It is not known whether the poor growth and inefficient utilization 
of feed result from poor digestion, poor absorption, or some other 
abnormality of metabolism. 

The present study was undertaken to determine the effect of 
dietary molybdenum upon voluntary feed intake, efficiency of feed 
utilization, digestibility of nutrients, and body weight changes of 
rats and rabbits. 

Experimental 

Weanling rats of the Long-Evans strain and weanling rabbits 
of the Dutch and New Zealand breeds were used as experimental 
animals. Rats were housed and fed individually in stainless steel 
cages with wire floors; rabbits were similarly fed in galvanized 
cages. The experimental diet for rats was prepared from purified 
ingredients and consisted of the following (in per cent): sucrose, 30; 
corn starch, 39; vitamin-free casein, 16; corn oil, 5; cellulose, 5; 
mineral mixture, 3; NaCl, 1; vitamin mixture, 1 (cellulose, "alpha- 
eel"; mineral mixture, USP XIV; vitamins, Vitamin Diet Fortifica- 
tion, Nutritional Biochemicals Corp, Cleveland, Ohio). Molybde- 
num was added as Na 2 Mo04.2H 2 in amounts to provide 500 and 



130 Quarterly Journal of the Florida Academy of Sciences 

1000 ppm molybdenum for two experimental diets. The control 
diet contained Na 2 C0 3 in amounts which provided an intake of 
sodium equal to that of the experimental diets. 

One group of rats (Table 1) was supplied the diets ad libitum 
and another group was pair-fed in order to equalize feed intake 
and provide a more critical measure of the effect of molybdenum 
upon feed efficiency. Those fed ad libitum were provided a 
weighed amount daily, and refused feed was weighed daily and 
discarded. Rats were randomly assigned to treatments on the 
basis of sex, litter, and body weight. Pair-fed animals were of the 
same sex, age, litter, and equal body weight. The rat of each 
pair consuming the molybdate diet was fed ad libitum, and the 
intake of the pair control was limited to that of the pair mate. 
Necessary precautions were taken to prevent feed wastage. Initial 
and weekly body weights were recorded. The feeding period for 
rats fed ad libitum was six weeks; four weeks for pair feeding. 

Molybdenum in the same form and the same concentration 
added to the diet was also supplied in the drinking water to eight 
rats which were fed control diets ad libitum. Daily intakes of the 
molybdate solution and of feed were recorded. 

At the end of the six-week feeding period, dietary treatments 
for rats fed ad libitum were reversed for one week and changes in 
feed intake and body weights were recorded. 

The diet for rabbits was a complete commercial pelleted feed 
(Purina Rabbit Checkers). Molybdenum as Na 2 Mo0 4 .2H 2 was 
added in amounts which provided 1000 and 2000 ppm of molyb- 
denum. Control diets contained Na 2 C0 3 in amounts which pro- 
vided sodium equal to that of the experimental rations. Eight- 
week-old Dutch and New Zealand rabbits were fed the molyb- 
date rations ad libitum, and in a second trial four-week-old Dutch 
rabbits were pair-fed for three weeks. 

Nutrient digestibility was determined during growth trials using 
the conventional digestion trial technique. Fecal collections were 
made for seven days, and dry matter and protein in feed and 
feces determined using methods outlined by A.O.A.C. (1960). Gross 
energy of feed and feces was determined with the adiabatic bomb 
calorimeter. Statistical calculations were based upon analysis of 
variance or Student's f for paired data (Snedecor, 1956). 



Arrington, Ammerman, Moore: Molybdenum Toxicity 131 

Results and Discussion 

Data representing feed intake, weight gains, and efficiency of 
feed utilization for rats are recorded in Table 1. One thousand 
ppm molybdenum either in the diet or in drinking water signifi- 
cantly reduced intake, gain, and feed to gain ratio of rats fed ad 
libitum, but five hundred ppm was not effective. When rats were 
pair-fed, both levels of molybdenum significantly reduced weight 
gain and efficiency of feed utilization for growth (Table 1). 

TABLE 1 
Feed intake, weight gain, and feed to gain ratio of rats fed molybdenum 







Av. daily 


Mo 


Av. daily 


Feed per 


H 2 




No. 


feed intake, 


intake, 


gain, 


unit gain, 


intake, 


Treatment 


rats 


gm 


mg/day 


gm 


gm 


ml/day 






Ad libitum feeding 








Control 


6 


12.2 


— 


2.8 


4.7 


22.9 


500 ppm Mo 














in feed 


6 


11.9 


6.0 


2.4 


5.1 


— 


1000 ppm Mo 














in feed 


6 


9.3* 


9,3 


1.1** 


8.6** 


20.6 


500 ppm Mo 














in water 


4 


12.8 


7.5 


2.6 


5.1 


15.0 


1000 ppm Mo 














in water 


4 


8.1** 


13,3 


0.4** 


37.6** 


13.4 






Pair feeding 








Control 


8 


11.4 


— 


3.2 


3.5 


— 


500 ppm Mo 


8 


11.4 


5.7 


2.7** 


4.2** 


— 


Control 


8 


7.7 





1.4 


5.7 


— 


1000 ppm Mo 


8 


7.7 


7.4 


0.8** 


10.8** 


— 



* Significantly different (P<0.05) from control within each column. 
** Significantly different (P<0.01) from control within each column. 



Rats consuming water containing 1000 ppm molybdenum con- 
sumed least feed, gained least, and were the most inefficient in 
utilization of the feed consumed. It may be noted that these rats 
consumed a greater amount of molybdenum than those provided 
molybdenum in the diet, although the total intake of water con- 
taining molybdenum was voluntarily restricted. Water intake of 



132 Quarterly Journal of the Florida Academy of Sciences 

control rats was not experimentally restricted, and the lower fluid 
intake by the rats consuming the molybdate solution may have 
accounted in part for the lower feed intake. 

Two thousand ppm molybdenum significantly reduced (P<0.01) 
feed intake and gain, but not feed efficiency of rabbits (Table 2). 





TABLE 2 

Feed intake, weight gain, 
gain ratio of rabbits fed 


and feed to 
molybdenum 




Treatment 


Av. daily 
feed intake 
(gm) 




Av. daily 
gain 
(gm) 




Feed per 

unit gain 

(gm) 


Control 
1000 ppm 
2000 ppm 


74 
67 

44** 




24 
22 
14** 




3.15 
3.16 
3.14 



** Significantly different (P<0.01) from control within each column. 

One thousand ppm in this natural diet appeared to reduce intake 
and gain, but the difference was not significant. When rabbits were 
pair-fed, no differences in weight gain or feed efficiency were ob- 
served. It appears that the effect of molybdenum upon rabbits 
under these conditions was only one of reducing feed consumption, 
and the different response of rats suggests a species difference. 
The feed intake per unit of body weight was greater for rats than 
for rabbits; thus equal concentration of molybdenum in the diet 
resulted in a greater intake of molybdenum per unit of body weight 
for rats. This may account in part for the apparent greater toler- 
ance of molybdenum by rabbits. 

Since a major effect of molybdenum appeared to be one of re- 
ducing feed intake, the average daily intakes of rats for the first 
seven days and the daily intake by weeks for the remaining five 
weeks are plotted in Fig. 1. Intakes of the molybdate rations and 
of the control rations with molybdate in the water were reduced 
in the first 24 hours and remained lower throughout the feeding 
period; however, 500 ppm caused only a slight reduction in intake. 
No gross abnormalities other than poor growth were observed 
during the relatively short experimental period. Animals appeared 
to be hungry although adequate molybdate diets were present. 



Arrington, Ammerman, Moore: Molybdenum Toxicity 



133 



-/h 





• CONTROL 
O 500 PPM DIET 
■x I 000 PPM DIET 



12 3 4 5 6 7 
DAYS 



y/- 



TREATMENTS 

CHANGED 



3 4 5 
WEEKS 



7 



Fig. 1. Feed consumption of rats fed molybdenum and changes in 
intake after reversing dietary treatments. 

When experimental animals which had been consuming the molyb- 
date diets for six weeks were changed to the control diet, there was 
a rapid increase in both intake and gain (Table 3). Rats which 
had been consuming the higher levels of molybdenum and grow- 

TABLE 3 

Changes in feed intake and weight change of rats 
after reversing dietary treatments 



Former Intake Change Change Weight 

intake Reversed 1st day 1st day week change 

gm/day gm/day % % gm/week 



Former 
diet 



500 ppm Mo 
1000 ppm Mo 
1000 ppm Mo 

in water 
Control 



11.7 
9.6 

8.7 

12.1 



Control 
Control 



13.5 
14.9 

14.6 



Dist H,0 
1000 ppm 

diet 10.7 



+ 15 

+55 

+68 
-12 



+ 10 
+51 

+53 

-17 



+23 

+37 

+45 
—20 



134 Quarterly Journal of the Florida Academy of Sciences 

ing least gave the greatest response in feed intake and gain. Rats 
changed from the control diet to molybdate diets or solution de- 
creased the intake of feed and lost weight within one week, but 
the decrease was not equivalent to increases made when changes 
were made from molybdenum to control diets. 

Digestibility of nutrients was not altered by molybdenum in 
either rats or rabbits in this study (Table 4). The average digest- 
ibility by rats consuming the purified diet was considerably greater 
than that by rabbits consuming the natural ration, but within each 
species, molybdenum was without effect. 

TABLE 4 

Digestion coefficients of nutrients by rats and rabbits fed molybdenum 



Treatment animals matter Protein Energy 

Rats 

Control 6 91.8 91.8 91.9 

500 ppm Mo— diet 6 91.5 90.8 91.9 

1000 ppm Mo— diet 6 91.8 91.1 92.2 

500 ppm Mo— water 4 92.8 92.4 92.8 

1000 ppm Mo— water 4 92.3 90.8 92.7 



Control 6 67.0 76.8 68.1 

1000 ppm Mo 4 66.7 72.4 67.2 

2000 ppm Mo 5 69.5 77.2 70.3 



The poor growth of experimental animals provided molyb- 
denum in the diet or in water is obviously accounted for, in part, 
by the reduced feed intake. Poor feed efficiency of rats fed ad 
libitum and lower gain and feed efficiency of rats which were 
equalized pair-fed suggested that there is an effect of molybdenum 
independent of nutrient intake. The absence of an effect upon 
nitrogen digestibility as reported by Johnson and Miller (1963) or 
upon digestibility of nutrients in this study suggests that the action 
of molybdenum occurs at a point in nutrient utilization other than 
in digestion. 



No. 


Dry 


animals 


matter 




Rats 


6 


91.8 


6 


91.5 


6 


91.8 


4 


92.8 


4 


92.3 




Rabbits 


6 


67.0 


4 


66.7 


5 


69.5 



Arlington, Ammerman, Moore: Molybdenum Toxicity 135 

Summary 

Molybdenum as NaMo04.2H 2 was fed to growing rats in a 
purified diet or in water and was added to a natural ration for 
rabbits. One thousand ppm dietary molybdenum significantly re- 
duced voluntary feed intake, growth, and efficiency of feed utili- 
zation of rats, but 500 ppm was ineffective. Molybdenum in the 
drinking water had a similar effect. When rats were equalized 
pair-fed, those consuming 500 or 1000 ppm molybdenum gained 
less than pair mates fed control diets and were thus less efficient 
in utilization of feed (P<0.01). Rabbits fed 2000 ppm molybde- 
num consumed less feed and gained less than controls (<0.01), but 
feed efficiency was not affected. One thousand ppm did not sig- 
nificantly reduce intake or gain of rabbits. Experimental rats 
changed to control diets, after consuming molybdate diets for six 
weeks, made immediate responses in terms of increased feed in- 
takes and weight gain. Digestibility of dry matter, protein, and 
energy was not affected by dietary molybdenum in either rats or 
rabbits. 

Acknowledgment 

This study supported in part by Moorman Manufacturing Co., 
Quincy, Illinois. The technical assistance of M. R. Ziegler is 
acknowledged. 

Literature Cited 

Association of Official Agricultural Chemists. 1960. Official and 
tentative methods of analysis. Association of Official Agricultural 
Chemists, Washington, D. C, 832 pp. 

Dick, A. T. 1956. Molybdenum in animal nutrition. Soil Sci., vol. 81, 
pp. 229-236. 

Johnson, H. L., and R. F. Miller. 1961. The interrelationships between 
dietary molybdenum, copper, sulfate, femur alkaline phosphatase ac- 
tivity and growth of the rat. Jour. Nutr., vol. 75, pp. 459-464. 

. 1963. Possible mechanisms for dietary molybdenum toxicity in the 

rat. Jour. Nutr., vol. 81, pp. 271-278. 

Miller, R. F., and R. W. Engel. 1963. Interrelationships of copper, mo- 
lybdenum and sulfate in nutrition. Federation Proc, vol. 19, pp. 666- 
677. 

Monty, K. J., and Ellen M. Click. 1961. A mechanism for the copper- 
molybdenum interrelationship. III. Rejection by the rat of molybdate- 
containing diets. Jour. Nutr., vol. 75, pp. 303-308. 



136 Quarterly Journal of the Florida Academy of Sciences 

Snedecor, G. W. 1956. Statistical methods. Iowa State College Press, 
Ames, Iowa, 534 pp. 

Underwood, E. J. 1962. Trace elements in human and animal nutrition. 
Academic Press, New York, N. Y., edition 2, 429 pp. 

Department of Animal Science, University of Florida, Gaines- 
ville, Florida. Florida Agricultural Experiment Stations Journal 
Series No. 1915. 



Quart. Jour. Florida Acad. Sci. 28(1) 1965 



FLORIDA ACADEMY OF SCIENCES 

Institutional Members for 1965 

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The next annual meeting will be at Florida Presbyterian Col- 
lege, St. Petersburg, March, 1966. 



FLORIDA ACADEMY OF SCIENCES 

Founded 1936 



OFFICERS FOR 1965 

President: O. E. Frye, Jr. 

Game and Fresh Water Fish Commission 

Tallahassee, Florida 

President Elect: Margaret Gilbert 

Department of Biology, Florida Southern College 

Lakeland, Florida 

Secretary: John D. Kilby 

Department of Zoology, University of Florida 

Gainesville, Florida 

Treasurer: John S. Ross 

Department of Physics, Rollins College 

Winter Park, Florida 

Editor: Pierce Brodkorb 

Department of Zoology, University of Florida 

Gainesville, Florida 



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Quarterly Journal 

of the 



fQFL 



Florida Academy of Sciences 



Vol. 28 June, 1965 No. 2 



CONTENTS 

Oxidation of organic sulfides by dinitrogen tetroxide 

Robert D. Whitaker and Carole L. Bennett 137 

The phyllosoma larvae of Parribacus Harold W. Sims, Jr. 142 

Ciguatera poisoning from barracuda Edward Larson and Luis R. Rivas 173 

Young common snook on the coast of Georgia 

Thomas L. Linton and William L. Rickards 185 

Food of Neoseps, the Florida sand skink 

Charles W. Myers and Sam R. Telford, Jr. 190 

Herring gulls diving for starfish 

Lowell P. Thomas and Shirley B. Thomas 195 

New taxa of fossil birds Pierce Brodkorb 197 

Ecology of the indigo bunting in Florida David W. Johnston 199 

Present status of the beaver in Florida 

James N. Layne and Bette S. Johns 212 

Dietary copper and enzymes in rabbit semen D. W. Stanley, 

R. L. Shirley, L. R. Arrington, and A. C. Warnick 221 

Florida Academy of Sciences award for 1965 225 

Florida Academy of Sciences charter and by-laws 226 



Mailed June 25, 1965 



Quarterly Journal of the Florida Academy of Sciences 
Editor: Pierce Brodkorb 



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QUARTERLY JOURNAL 

of the 

FLORIDA ACADEMY OF SCIENCES 



Vol. 28 June, 1965 No. 2 



OXIDATION OF ORGANIC SULFIDES BY DINITROGEN 

TETROXIDE 

Robert D. Whitaker and Carole L. Bennett 

The physical properties of dinitrogen tetroxide (m.p. — 11.5°, 
b.p. 21.3°), its high solubility in common solvents such as chloro- 
form and carbon tetrachloride, and its ready commercial avail- 
ability make it a most convenient oxidizing agent. It is particu- 
larly useful for the oxidation of organic sulfides to the correspond- 
ing sulfoxides (Addison and Sheldon, 1956; Whitaker and Sisler, 
1960). 

R 2 S + N 2 4 -> RoSO + N 2 3 

In most cases, the sulfoxide is produced in quantitative yield. 
There is no danger of sulfone contamination since dinitrogen 
tetroxide will not oxidize sulfoxides to sulfones. Thus, unlike 
oxidations using H 2 2 or Cr0 3 where the control of stoichiometry 
is critical, excess dinitrogen tetroxide may be used. 

Dinitrogen tetroxide and sulfoxides form molecular addition 
compounds of the Lewis acid-base type (Addison and Sheldon, 
1956), and it has been suggested that the formation of these ad- 
ducts inhibits the further oxidation of the sulfoxide. In an attempt 
to test this hypothesis, we have studied the reaction between di- 
nitrogen tetroxide and bis(chloromethyl) sulfide. We felt that the 
relatively electronegative chlorine atoms would significantly de- 
crease the electron density of the sulfur atom. The result might 
then be that Lewis acid-base interaction between any bis(chloro- 
methyl) sulfoxide produced initially and the dinitrogen tetroxide 
would be greatly weakened or eliminated. In this way, we thought 
that possibly the sulfone would be produced or perhaps some en- 
tirely different product. 



ihstitwioh JUt * 1965 



138 Quarterly Journal of the Florida Academy of Sciences 

This work was done while the junior author was a National 
Science Foundation Undergraduate Research Participant. Some 
of the results were presented to the Organic Division at the 1964 
Meeting-in-Miniature of the Florida Section, American Chemical 
Society. 

To our surprise, when bis(chloromethyl) sulfide was allowed 
to react with dinitrogen tetroxide, the subsequent removal of excess 
dinitrogen tetroxide and nitrogen trioxide invariably led to an ex- 
plosion. On several occasions, its violence was sufficient to shat- 
ter the glass apparatus in which the reaction was run. We have 
cautiously investigated this reaction in some detail since no pre- 
vious oxidation of an organic sulfide with dinitrogen tetroxide had 
led to such a violent reaction. The results of our investigation are 
summarized in Table 1. 

We have found that dinitrogen trioxide (probably reacting 
through the NO radical) attacks unreacted bis(chloromethyl) sul- 
fide to give the violent, highly exothermic reaction. Apparently, 
bis(chloromethyl) sulfoxide and dinitrogen trioxide are formed ini- 
tially when bis(chloromethyl) sulfide reacts with dinitrogen tetrox- 
ide at 0° (Exp. 1, 2, Table 1). Subsequent warming of the re- 
action mixture plus removal of excess dinitrogen tetroxide, which 
serves to concentrate the solution of unreacted bis(chloromethyl) 
sulfide and dinitrogen trioxide, causes the reaction between these 
species to become very rapid. Direct reaction between dinitrogen 
trioxide and bis(chloromethyl) sulfide likewise leads to a violent 
reaction (Exp. 3, Table 1). When the reaction is run under condi- 
tions which inhibit the formation of dinitrogen trioxide (Exp. 9, 
Table 1), the only product is bis(chloromethyl) sulfoxide, and the ex- 
cess dinitrogen tetroxide can be safely removed at room tempera- 
ture. Since it has been shown that nitric oxide will not react with 
unsubstituted alkyl sulfides (Longhi, 1963), it is clear that the pres- 
ence of the chlorine atoms in the sulfide molecule leads to this 
novel reaction. The possibility that the sulfoxide or sulfone is 
initially produced and then reacts with one of the nitrogen oxides 
is ruled out by our results (Exps. 4, 5, 6, 7, Table 1). In addition, 
no trace of sulfone was ever detected in the reaction mixtures. 
It is not possible to decide whether the attack of the dinitrogen 
trioxide (or nitric oxide) occurs principally at the Cl-C bond or 
at the C-S bond of the bis(chloromethyl) sulfide. Since SC1 2 is one 



Whitaker and Bennett: Dinitrogen Tetr oxide 



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